ThesisPDF Available

Plant pollinator network of Kashmir valley

Authors:

Abstract

Understanding the structure and function of plant-pollinator networks has emerged as a priority research area, and natural history observations of the relationship between plants and their pollinators enable us to reconstruct these networks. Here we report the structure of a plant-pollinator network from Kashmir Himalaya, including its modularity, integration of alien species, and the effects of extinction simulation. We recorded the network by using natural history observations to document the frequency of pollinators on plants. We analyzed data for network visualization, indices calculation, null model comparisons and extinction simulation. A total of 230 plant and 80 pollinator species formed 1958 (11% of the possible) interactions in the network. Among the plants Malus domestica and among the pollinators Apis mellifera and A. cerana formed the largest number of interactions with significant (p<0.004) interaction patterns. Network cumulative degree distribution depicted a higher number of degree levels in pollinators than plants (p<0.001). Nestedness, connectance and cluster coefficient revealed moderately high number of realized interactions, thereby indicating greater structural and functional stability in the network. Eight strongly defined modules (p<0.001) were observed in the network. The Ephedra module exclusively comprised of native species whereas the Apis module comprised all the four different types of interacting species (native and non-native plants and pollinators) integrated highest number of non-native species. Interestingly, in the network 40% of interactions were by non-native species reflecting well integration. Extinction simulations suggested that the network would collapse more quickly when the most connected pollinators are removed, rather than the most connected plant species. Our study is the first assessment of structure and function of a plant pollinator network from the Himalayan biodiversity hotspot; and looking ahead, we discuss the ecological and economic implications of such interactions in an era of global biodiversity crisis.
Studies on Plant and Insect Visitor
Interactions in Kashmir Valley
DISSERTATION
Submitted to the University of Kashmir for the
Award of Degree of
MASTER OF PHILOSOPHY (M. Phil.)
IN
BOTANY
By
Zubair Ahmad Rather
(M.Sc. & NET)
Under the Joint Supervision of
Dr. Shoukat Ara
(Co-supervisor)
Dr. Anzar A. Khuroo
(Supervisor)
CENTRE FOR BIODIVERSITY & TAXONOMY
DEPARTMENT OF BOTANY
FACULTY OF BIOLOGICAL SCIENCES
University of Kashmir, Srinagar-190006
(NAAC Accredited Grade ‘A’)
2016
Certified that the d
issertation entitled
Interactions in Kashmir Valley
Hazratbal, Srinagar
Botany, embodies original research work carried out by
Rather under our
joint supervision. This work has not been submitted in part
or in f
ull for this or any other degree before.
It is further certified that
Department and
fulfills
Botany.
Dr. Shoukat Ara
(Co-supervisor)
Associate Professor
Division of Environmental Sciences
SKUAST-K, Shalimar
Srinagar, J&K
DEPARTMENT OF BOTANY
UNIVERSITY OF KASHMIR
HAZRATBAL, Srinagar, Kashmir
No……............
Dated…………..
Certificate
issertation entitled
Studies on Plant and Insect Visitor
Interactions in Kashmir Valley
submitted to
the University of Kashmir,
for award of the degree of "
Master of Philosophy
Botany, embodies original research work carried out by
Mr. Zubair Ahmad
joint supervision. This work has not been submitted in part
ull for this or any other degree before.
It is further certified that
the candidate
has put in the required attendance in the
fulfills
all the requirements for the
award of M.
Division of Environmental Sciences
Dr. Anzar A. Khuroo
(
Supervisor
Senior Assistant Professor
Department of Botany
University of Kashmir
Srinagar, J&
Head
Department of Botany
University of Kashmir
Srinagar-190006
DEPARTMENT OF BOTANY
UNIVERSITY OF KASHMIR
HAZRATBAL, Srinagar, Kashmir
-190006
No……............
...
Dated…………..
Studies on Plant and Insect Visitor
the University of Kashmir,
Master of Philosophy
" in
Mr. Zubair Ahmad
joint supervision. This work has not been submitted in part
has put in the required attendance in the
award of M.
Phil. degree in
Dr. Anzar A. Khuroo
Supervisor
)
Senior Assistant Professor
Department of Botany
University of Kashmir
Srinagar, J&
K
Affectionately
Dedicated
To
My Loving Parents
Caring Wife
and
Dear Daughter
Acknowledgements
I am highly grateful to Almighty Allah for His each and every
blessing.
I am immensely thankful and indebted to my supervisor Dr. Anzar A.
Khuroo, Senior Assistant Professor, Department of Botany,
University of Kashmir and co-supervisor, Dr. Shoukat Ara, Senior
Associate Professor, Division of Environmental sciences, SKUAST-
Kashmir for their invaluable suggestions, timely help and dexterous
supervision in cooperative and sympathetic approach at all stages of
this research study.
I am grateful to Head Department of Botany, Professor Inat-ul-lah
Tahir for providing necessary support, facilities and conductive
atmosphere during the course of this study.
I am highly thankful to Indian Council of Agricultural research
(ICAR) for providing me the financial assistance under the project
Pollination management research in apples and other fruits in
Kashmir valleyNiche Area of Excellence (NAE-II) project no. F. No.
10(5)/2012-EPD dated: 26-03-2012.
I am also highly thankful to Dr. Manzoor Ahmad Paray, Senior
Associate Professor, Division of Entomology, SKUAST- Kashmir, Dr.
Sajad Hussain Parey, Young Scientist DST, Dr. Munaza Yaqoob, Dr.
Rizwana Khursheed, Dr. G. H. Rather, Dr. Barkat Hussain, and
entire faculty working at Research and training centre for pollinators
pollinizers and pollination management for their help in surveys and
insect visitor identification.
I am thankful to all my esteemed teachers for their encouragement
especially Shameem Ahmad Rather for his moral, emotional and
economical help, Dr. Rifat H. Raina for helping in identification of
insect visitors, Akhtar H. Malik for identification of pollen and nectar
producing plant species and entire staff of Centre for Biodiversity and
Taxonomy for their necessary support.
I owe sincere thanks to my lab mates Waseem Ahmad Bhat, Miss
Ferhana Majid, Miss Gousia Mehraj, Miss Shugufta Raheed, Mr.
Maroof Hamid, Mr. Rameez Ahmad and Mr. Marifatul-Haq for
their help and encouragement during the entire course of this study.
I have no words to express the immense gratitude, affection and
admiration for my family especially my dear parents (Mr. Ghulam
Nabi Rather and Hajra bejum) for their unconditional love, affection,
blessings, encouragement, great moral and emotional support. I will
never forgrt their hardship in growing me up and their blind faith in
me. I am greatly indebted to my caring wife (Mrs. Bilqees) for her
inculcable help, valuable suggestions and unconditional love during all
the time.
Zubair Ahmad Rather
CONTENTS
Page Nos.
Summary 1-8
Chapter 1
Taxonomic diversity of foraging plants and their
insect visitors 9-57
Introduction
Review of literature
Study Area
Materials and Methods
Results
Discussion
Conclusions
References
9
10
13
14
16
43
46
47
Chapter 2
Diversity in floral traits with reference to insect
visitors 58-110
Introduction
Review of literature
Materials and Methods
Results
Discussion
Conclusions
References
58
59
66
70
95
100
101
Chapter 3
Determination of pollinator effectiveness
among insect
visitors 111-150
Introduction
Review of literature
Materials and Methods
Results
Discussion
Conclusions
References
111
112
117
118
139
142
143
Chapter 4
Characterization of generalized and specialized plant
insect interactions 151-201
Introduction
Review of literature
Materials and Methods
Results
Discussion
Conclusion
References
151
152
154
155
191
194
196
LIST OF TABLES
Table No.
Title of Table Page No.
1.1
Diversity of foraging plants, their family, English name,
Kashmiri name, habit, purpose of cultivation, insect visitor (s),
reward, geo-coordinates and site of record in Kashmir Himalaya.
18
1.2
Taxonomic conspectus of insect flower visitors recorded during
the present study in the Kashmir Himalaya.
38
2.1 Categories of floral display size. 67
2.2 Categories of inflorescence display size. 68
2.3 Floral
trait database recorded in the pollinator flora of Kashmir
Himalaya.
79
3.1
Pollen deposition on different plant species by different insect
visitors.
135
3.2
Visitation rate of different flower visitors on different plant
species.
137
4.1 Inventory of plant pollinator generalization and specialization. 166
LIST OF FIGURES
Fig. No. Title of Figure Page No.
1.1 Location map of the study area 13
1.2 Number
of species distributed in different families of the
flowering plants recorded during present study.
41
1.3 Percentage contribution of different growth-
forms in the
flowering plants recorded during present study.
41
1.4 Wild and cultivated status of the
flowering plants recorded
during present study (number and percentage)
42
1.5
Number and percentage of the flowering plants with reference
to insect visitor reward (PL=pollen, NC=nectar, PN= both
pollen and nectar)
42
1.6 Number of species belonging to
different families of insect
flower visitors recorded during present study.
43
2.1
Contribution of different inflorescence types in the pollinator
flora.
71
2.2 Number of species in different floral display size classes 72
2.3 Number of species
belonging to different inflorescence display
size classes
72
2.4
Distribution of flower reward type in terms of number of
species and percentage among pollinator flora
73
2.5
Number of species belonging to different floral shapes recorded
in the pollinator flora of Kashmir Himalaya
74
2.6 Distribution of
flower symmetry in terms of number of species
and percentage among the pollinator flora
74
2.7
Distribution of floral sex in terms of species richness and
percentage among the pollinator flora.
75
2.8 Distribution of
flower chromatism in terms of number of
species and percentage among the pollinator flora.
75
2.9
Number of species belonging to different corolla colour types in
the pollinator flora of Kashmir Himalaya.
76
2.10 Number of species
belonging to different anther colour types in
the pollinator flora of Kashmir Himalaya.
77
2.11
Number of species belonging to different pollen colour types in
the pollinator flora of Kashmir Himalaya.
78
2.12 Presence and absence of flower scent among
the pollinator
flora.
78
3.1
Pollen deposition and visitation rate of different flower visitors
on Abelia grandiflora.
119
3.2
Pollen deposition and visitation rate of different flower visitors
on Actinidia deliciosa.
120
3.3
Pollen deposition and visitation rate of different flower visitors
on Aesculus indica.
120
3.4
Pollen deposition and visitation rate of different flower visitors
on Alcea rosea.
121
3.5
Pollen deposition and visitation rate of different flower visitors
on Antirrhinum majus.
121
3.6
Pollen deposition and visitation rate of different flower visitors
on Berberis lycium.
122
3.7
Pollen deposition and visitation rate of different flower visitors
on Brassica compestris.
122
3.8
Pollen deposition and visitation rate of different flower visitors
on Brassica rapa.
123
3.9
Pollen deposition and visitation rate of different flower visitors
on Cichorium intybus.
123
3.10
Pollen deposition and visitation rate of different flower visitors
on Colchicum luteum.
124
3.11
Pollen deposition and visitation rate of different flower visitors
on Convolvulus arvense.
124
3.12
Pollen deposition and visitation rate of different flower visitors
on Coriandrum sativum.
125
3.13
Pollen deposition and visitation rate of different flower visitors
on Cydonia oblonga.
125
3.14
Pollen deposition and visitation rate of different flower visitors
on Epilobium hirsutum.
126
3.15
Pollen deposition and visitation rate of different flower visitors
on Hypericum hookerianum.
126
3.16
Pollen deposition and visitation rate of different flower visitors
on Hypericum perforatum.
127
3.17
Pollen deposition and visitation rate of different flower visitors
on Lavatera cashemiriana.
127
3.18
Pollen deposition and visitation rate of different flower visitors
on Mahonia borealis.
128
3.19
Pollen deposition and visitation rate of different flower visitors
on Malus domestica.
128
3.20
Pollen deposition and visitation rate of different flower visitors
on Prunus armeniaca.
129
3.21
Pollen deposition and visitation rate of different flower visitors
on Punica granatum.
129
3.22
Pollen deposition and visitation rate of different flower visitors
on Rosa webbiana.
130
3.23
Pollen deposition and visitation rate of different flower visitors
on Rosa indica.
130
3.24
Pollen deposition and visitation rate of different flower visitors
on Sisymbrium irio.
131
3.25
Pollen deposition and visitation rate of different flower visitors
on Sternbergia vernalis.
131
3.26 Pollen depositions by Apis mellifera and A. cerana
on different
plants.
132
3.27 Visitation rate of Apis mellifera and A. cerana
on different
flowering plants.
133
3.28 Pollen depositions by Xylocopa valga, Apis mellifera and
A.
cerana on different flowering plants.
134
3.29 Visitation rate of Xylocopa valga on different flowering plants. 134
LIST OF PLATES
Plate No.
Title of Plate Following
Page No.
1.1 Collection of insect visitors. 15
1.2
Representative pollen and nectar producing plant
species.
16
1.3
Representative pollen and nectar producing plant
species.
16
1.4 Representative pollen and nectar producing
plant
species.
16
1.5
Representative pollen and nectar producing plant
species.
16
1.6 Representative insect visitors. 17
1.7 Representative insect visitors. 17
2.1 Floral
traits in pollen and nectar producing plant
species.
70
2.2 Floral traits
in pollen and nectar producing plant
species.
70
2.3 Floral
traits in pollen and nectar producing plant
species.
70
3.1
Representative pollen deposited stigmas after single
visit deposition.
118
3.2 Representative pollen deposited stigmas after
single
visit deposition.
118
4.1 Generalization in Dahlia daenranthema. 155
4.2 Generalization in different foraging plant species. 155
4.3 Generalization in different foraging plant species. 155
4.4 Specialization in different foraging plant species. 155
Studies on Plant and Insect Visitor
Interactions in Kashmir Valley
Studies on Plant and Insect Visitor
Interactions in Kashmir Valley
Studies on Plant and Insect Visitor
Interactions in Kashmir Valley
Summary
1
SUMMARY
In nature, living organisms never live in isolation. The diverse kinds of
plants, animals, microbes and other life forms - known as biodiversity - always
interact with each other to constitute biological communities, which in turn
interact with physical environment to form diverse ecosystems. Such a rich
biodiversity and the ecosystem they form, provide precious economical goods
and ecosystem services which are crucial for human survival. In fact
maintainance and sustainance of biodiversity depends on a complex set of
biological interactions, and therefore the scientific understanding of these
interactions assumes immediate relevance in the conservation and sustainable
use of biodiversity (Jordano et al., 2003; Campbell et al., 2011). Of the various
biological interactions existing in an ecosystem, interaction between flower of
plants and their insect visitors or flower-insect interaction is an important co-
evolutionary phenomenon that sustains a series of ecological and evolutionary
processes and ultimately underpins the overall maintenance of biodiversity and
ecosystem services (Proctor et al., 1996; Waser et al., 1996; Thompson, 1998).
Flower insect interaction is the most precious one, which is directly or
indirectly related to apiculture, pollination, honey, wax, royal jelly, fruit, seed
production and a lot of allied benefits to human society. Flower-insect
interaction is a mutualistic approach of interaction, each interacting life form
benefiting by the presence of other. This interaction involves the interaction
between two different life forms; flower of plants and their insect visitors and
may be opportunistic (generalized) and obligatory (specialized) (Frame, 2003).
Few flower-pollinator interactions are absolutely obligatory (Kearns, 1998)
where as most are generalized (Armbruster and Baldwin, 1998). Global
estimate of species diversity within angiosperms revealed composition of
250,000 plant species, among which 90% are mostly pollinated by animals
particularly insects.
In recent times the decline in managed and natural pollinators from last
Summary
2
several decades has resulted in negative consequences for wild and cultural
ecosystem pollination services. Indiscrimnate and non-judicial use of
pesticides (insecticide and herbicide) which is the direct outcome of
agricultural intensification has resulted in on-spot death of pollinators through
intoxification and is a major challenging threat to pollinators. Extinction of
some pollinators by different anthropogenic activities such as habitat
fragmentation, agricultural intensification, grazing, pesticide usage and
pollution of environment, climate change has led to parallel decline of their
interacting web members. Decline of flower-pollinator web has concerned
conservationists, orchardists and general public due to deteriorating quality and
quantity of both natural as well as cultural ecosystems.
The Himalaya represents one of the world’s tallest ranges of mountains,
which are located in the Indian subcontinent. Being one of the global
biodiversity hotspots, the Himalaya possesses a precious biological wealth of
great economic value and scientific interest. Kashmir Himalaya, lying in the
north-western part of Himalyas, is renowned for its natural landscape
characterized by a diverse and endemic biodiversity (Lawrence, 1895; Dar et
al., 2013). Notwithstanding rich biodiversity, very few research studies have
been carried out to explore the diversity of flowering plants and their insect
visitors in this Himalayan region (Williams, 1991; Saini et al., 2012; Ganie et
al., 2013; Paray et al., 2014). These studies have been either narrow in scope,
focussed on insect visitors only, without documenting the diversity of
flowering plants in detail or specific to cultivated plants such as apple. It is in
this context that the present study, using an integrated approach, was
undertaken to study the plant and insect visitor interactions in Kashmir Valley
with following objectives in mind:
Taxonomic diversity of foraging plants and their insect visitors
Diversity of floral traits with reference to insect visitors.
Determination of pollinator effectiveness among insect visitors
Characterization of generalized and specialized plant insect
interactions
Summary
3
Based on the results obtained during the present study, a total 227 plant
species belonging to 182 genera and 58 families were recorded, which serve as
foraging plants for different flower visitors. The annual herbaceous plants were
found as abundant source of pollen and nectar for flower visitors followed by
perennial herbs, perennial shrubs, trees and finally biennial herbs. Asteraceae
was found to be dominant family of pollen and nectar producing plants in
Kashmir Himalaya. The wild flora was found to be dominant source of pollen
and nectar. Majority of the pollen and nectar producing plants were found to
act as the source of both pollen and nectar (bi-dimensional) as compared to
pollen or nectar only (uni-dimensional). The study documented 70 flower
visitors belonging to 43 genera and 23 families which were found visiting 227
plants species. Family Apidae was the dominant group of flower visitors.
The present study revealed that there is a great diversity within the floral
traits that a flower displays for the attraction of pollinators. The attracting floral
traits guide and direct the flower visitor towards the ultimate reward which
serves as energy source for pollinators. Floral traits include both traits of an
inflorescence and its flowers. The study of inflorescence architecture of
foraging flora showed a great diversity with majority of plant species with
raceme inflorescence followed by cymose, capitulum, catkin etc. Floral display
size, the most important driver of pollinator attraction, showed a great
diversity. It was categorized into 5 classes each with specific range of display
size such as very small floral display (VSFD), small floral display (SFD),
medium floral display (MFD), large floral display (LFD), very large floral
display (VLFD) and great relation between floral display size and pollinator
visitation rate was observed, greater the display size greater the pollinator
visitation rate and vice versa. Infloresecnec display was also categorized into
five categories such as VSID, SID, MID, LID and VLID and among these
majorities of pollinator flora had very small inflorescence display (VSID). As
compared to single flower, inflorescence attracts the more number of flower
Summary
4
visitors due to increased inflorescence display size. Majority of pollen and
nectar producing plant species were bisexual serving source for both pollen and
nectar and a few were unisexual.
Further, flower shape is an important floral trait which is a main factor
in determining the foraging behaviour. A great diversity in flower shape was
observed and majority of pollinator flora had rotate shaped flowers. Analysis of
flower symmetry of pollinator flora revealed that majority of plants had
actinomorphic flowers serving as generalized towards pollinators. Generally,
two flower shapes were observed open access shapes include rotate, saucer,
bowl, cruciform, rosaceous and closed access flower includes bilabiate,
campanulate, trumpet, paplionaceous and caryphylaceous, personate. These
two shaped flowers have different interacting efficacies with pollinators and
thus modify generalization-specialization continuum range of interaction. It
was found that generally open access flowers acts as generalists towards their
pollinators as compared to closed accessed one. The results revealed that
symmetrical flowers act as generalized source and have more pollinator
visitation rate as compared to asymmetrical flowers. Majority of flowers were
found to show monochromatism as a floral trait followed by dichromatism and
multichromatism. Flower colour too had a great diversity and majority of
pollinator flora had yellow coloured flowers. Anther and pollen colour were
also found to show diversity among documented pollen and nectar producing
plants and majority of pollinator flora displays yellow colured anthers and
pollens, respectively. Floral scent presentation among studied pollinator flora
revealed that majority of plant species were without scent.
The present study characterized the pollen depositing ability and
visitation rate of eight (08) different insect visitor species on twenty five (25)
foraging plant species and differentiated efficient pollinators from inefficient
ones. e.g. Xylocopa valga from Eristalis tenax and Vanessa cashmeriansis. The
results reveal that different plant species have specific insect visitors, such as
Summary
5
Xylocopa valga deposits highest number (24) of pollens on the virgin stigma of
Abelia grandiflora followed by Bombus tunicatus (16), Apis mellifera (13),
Apis cerana (07) and visitation rate was found highest for Apis mellifera (04
visits/10 minutes) followed by Xylocopa valga on Abelia grandiflora flowers.
Likewise, Apis mellifera deposited highest (23) number of pollens followed by
Apis cerana (18) and Eristalis tenax (4) on virgin stigmas of Berberis lycium.
The highest visitation rate was recorded for Apis cerana (4) followed by other
three taxa (03 each) on Berberis lycium flowers. Apis mellifera deposited
highest (15) number of pollens followed by Apis cerana (11), Andrena flavipes
(8), Eristalis tenax (5) and Lasioglossum margenatum (4) on virgin stigmas of
Brassica compestris. Lasioglossum margenatum has highest visitation rate (06)
followed by Apis cerana (04), Apis mellifera (03), Andrena flavipes (02) and
(01) of Eristalis tenax on Brassica compestris flowers. Apis mellifera deposited
highest (15) number of pollens followed by Apis cerana (9), Andrena flavipes
(5) and Lasioglossum margenatum (3) on the virgin stigmas of Brassica rapa.
Visitation rate (5 visits/10 minutes) was recorded highest for Lasioglossum
margenatum and Apis cerana followed by Apis mellifera (4) and (03) Andrena
flavipes on Brassica rapa flowers.
Among the Apis genus, A. mellifera had more pollinator effectiveness as
compared to A. cerana both in terms of both pollen deposition as well as
visitation rate on Actinidia deliciosa, Alcea rosea, Cydonia oblonga, Cichorium
intybus, Convolvulus arvense , Epilobium hirsutum, Hypericum hookerianum,
H. perforatum, Lavatera cashmiriana, Mahonia borealis, Punica granatum,
Rosa indica, Sisymbrium irio, Sternbergia vernalis. It was found that Apis
mellifera had high pollen deposition ability than A. cerana and A. cerana had
highest visitation rate than A. mellifera on Aesculus indica, Berberis lyceum,
Brassica compestris, B. rapa, Colchicum luteum, Coriandrum sativum, Malus
domestica, Prunus armeniaca, Rosa webbiana. A comparative analysis of
pollen deposition by Apis mellifera and Apis cerana on different flowering
Summary
6
plants revealed that Apis mellifera deposited highest number of pollens than A.
cerana on all plant species. Xylocopa valga deposited the highest number of
pollens on virgin stigmas followed by Apis mellifera and A. cerana. Thus
Xylocopa valga serves as best pollen depositing insect visitor followed by
Bombus tunicatus, Apis mellifera, A. cerana, Andrena flavipes, Lasioglossum
marginbatum and finally Eristalis tenax.
The present study revealed that among the 215 plant pollinator
interactions 134 (62%) are generalized whereas 81 (38%) are specialized.
These two sets of interactions involve 215 foraging plant species and 70
different flower visitor species. Among 215 different flowering plant species
03 plant species are foraged by a single flower visitor species, 79 plant species
foraged by 2 species of flower visitors, 22 plant species by 3 flower visitor
species, 20 plant species by 4 flower visitor species, 11 plant species by 5
flower visitor species, 11 plant species by 6 flower visitor species etc. Plant
generalization specialization continuum revealed that Malus domestica and
Prunus domestica are foraged by the maximum number (23) of flower visitor
species belonging to 12 genera followed by Taraxacum officinale are foraged
by 21 species belonging to 12 genera of insect visitors. Pyrus communis and
Pyrus pyrifolia are foraged by 22 species belonging to 11 genera of flower
visitors. Pinus wallichiana was foraged by Bombus tunicatus and Bombus
trifasciatus, Aster thomsonii was foraged by Lassioglossum margenatum,
Lassioglossum himalayense, and Galinsoga parviflora by Episyrphus
balteatus, Frankliniella sp. Oxytropis cashmeriana is foraged by
Macroglossum nycteris only, Datura stramonium and Bidens tripartita by
Episyrphus balteatus only. 75 plant species are foraged only by Apis cerana
and A. mellifera.
Furthurmore Apis cerana is the most generalist pollinator visitor
foraging 205 plant species followed by Apis mellifera foraging 204 plant
species Eristalis tenax (41), Bombus trifasciatus (38), Bombus tunicatus (37),
Summary
7
Lasioglossum margenatum (35), Lassioglossum himalayense (30),
Lassioglossum nursei (29), and Episyrphus balteatus, Lassioglossum
rugolatum, Lassioglossum polyctor, Lasioglossum sublaterale, Lasioglossum
leucozonium by 26, Pieris brassicae forages 25 plant species, Xylocopa valga,
Xylocopa violacea forages 24 plant species, Formica sp., Andrena flavipes
forages 22 plant species, Vanessa cashmiriensis forages 21 plant species,
Bombus simillmus, Eoseristalis cerealis forages 19 plant species, Vanessa
cardui forages 14 plant species, Scathophaga stercoraria forages 13 plant
species, Bombus rufofasciatus, Bombus pyrosoma forages 12 plant species,
Bombus asiaticus, Eristalinus taeniops and Megachile rotundata forages 10
plant species, Bombus miniatus forages 9 plant species, Andrena patella,
Andrena floridula forages 8 plant species, Ceratina hieroglyphica, Hesperia
comma forages 7 plant species, Frankliniella sp., Sphaerophoria bengalensis,
Bibio marci, Plecia nearctica, Cetonia aurata forages 6 plant species,
Eristalinus aeneu, Metasyrphus bucculatus, Musca domestica, Polyommatus
icarus, Danais chrysippus forages 5 plant species, Sphaerophoria macrogaster,
Issoria lathonia forages 4 plant species, Mordella sp., Amegilla fallax, Vespa
velutina, Pyrrhocorus sp., Colias philodice, Pontia daplidice forages 3 plant
species, Ceratina propinqua, Sphecodes lasimensis, Sphecodes tantalus,
Coccinella magnifica, Mylabris pustulata, Scolia nobilitata, Halictus
constrictus, Colias fieldi, Macroglossum nycteris forages 2 plant species,
Macroglossum nycteris, Heriades sp., Scolia hauseri, Scolia soror, Scoliinae
proscoliinae, Spilomyia sulphurea forages only one plant species. The present
study revealed that 130 interactions were existing between generalized plant
species and generalized flower visitor species. 82 interactions were existing
between specialized plant and generalized flower visitor. Inaddition, 6
interactions were existing between generalized plant species and specialized
flower visitor and none interaction were existing between specialized plant
species and specialized flower visitor species.
Summary
8
Inconclusion, the results obtained during the present study can provide
a scientific platform for future research in the field of plant pollinator
interaction, pollinator food, pollen and nectar producing plants, pollination
biology, organismic interaction, diversity and co-extinction of biodiversity in
this Himalayan region. Cataloguing the diversity of pollen and nectar
producing plants enriches our knowledge about diversity and durability of
pollinator food resources, pollinator fauna wealth, magnitude and width of their
interacting web, its role and the need for conservation of flower pollinator web.
Besides documenting the food status of pollinators employed for natural as
well as agro-ecosystems, the diversity of pollen and nectar producing plants
reflects diversity of interacting pollinators and manifests in the form of most
important ecosystem service of pollination, which is crucial for maintenance
and sustenance of global biodiversity. The results can provide baseline
biodiversity information framework for conservation of plant insect interaction
and can help to cope with currently pressing problem of pollinator decline,
which has huge implications for production of different fruit crops of Kashmir,
and will help in overall maintenance and sustenance of biodiversity in Kashmir
Himalaya.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
9
INTRODUCTION
The huge diversity seen in the flowering plants, in addition to other
factors, is attributed to the interactions between flowers and insects that visit
them (Burkle, 2011). Insects which visit the flowers for their food are known
as flower visitors or insect visitors, and the plants visited by these flower
visitors are known as foraging plants (Kevan et al., 1983). In other words, the
plant species that support several insect visitors are potentially valuable forage
plants (Comba et al., 1999). The foraging plants provide pollen and nectar as
main rewards in the form of food to flower visitors and flower visitors act as
pollen carriers for the successful pollination and reproduction, thus help in
sustaining the plant life. Such a sort of socialistic mutualism is particularly
crucial in the maintenance of natural as well as cultural ecosystems, and the
wealth of economic goods and ecosystem services they provide to humankind
(Araujo, 2005).
Pollen and nectar producing plants are the nesting, feeding and mating
sites for flower visitors - commonly referred to as bee or forage flora. These
plant species are visited by bees and other pollinators for floral reward, such as
pollen and nectar for their survival and results into ecosystem goods and
services in the form of pollination, honey, wax, royal jelly, propolis (resinous
mixture collected from plants used for sealing the bee hive) and bee venom
production. The pollen and nectar, acts as dietary substance for pollinators
containing proteins, vitamins, minerals, carotenoids, fatty acids etc., fullfills
nutrient requirements of pollinator. In fact, the bee- keeping an agro-
horticultural forest-based industry is the reflection of floral resources present in
a geographical region. Both quantity and quality of bee-keeping of a region
directly depends on the diversity, abundance and durability of floral resource
they utilise. Honey quality is assessed by the flora that bees utilise as forage
plants, thus supporting income generation at global scale.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
10
REVIEW OF LITERATURE
Research investigations relating to taxonomic diversity of foraging or
bee flora and their insect visitors have been carried out in different parts of the
world. The bee flora is defined as the number of plant species visited by bees
for pollen and nectar for their survival and production of ecosystemic outcome
in the form of honey and providing ecological service pollination (Pereira,
1990; Sakuragui, 2011). It has been suggested that the identification of bee
flora of a region can optimise honey production by using appropriate source of
forage plants (Sakuragui, 2011).
Crane et al. (1984) identified the foraging plants or bee flora on the
basis of foraging behaviour of European bee and Africanized bee (the latter is a
hybrid between European and African honey bee). Partap (1997) found that
bee flora serves as an important natural input in bee-keeping and best source of
pollen and nectar for pollinators, and emphasized that the floral resource
diversity interacting with pollinator diversity underpins the pollination and
bee-keeping. Ramalho et al. (1990) reported 288 plant species as pollen and
nectar producing plants for pollinators, and among these 80 plant species were
found important to stingless bees and “Africanized” honey bees in the
Brazilian Neo-tropics. Momose et al. (1998) studied pollinator foraging in
Dipterocarp forest of Malaysia and found 270 plant species as foraging plants
to different types of pollinators. They also found 12 categories of pollination
system and reported 32% plants were foraged and pollinated by social bees
followed by beetles with 20%. Ebenezer and Olugbenga (2010) found 26
species as potential foraging plants by pollinators from the Savanna zone in
Guinea. Abdullahi et al. (2011) found 103 plant species that were potential
foraging plants for pollinators (honeybees) from Nigeria. Sakuragui et al.
(2011) identified 135 plant species belonging to 104 genera and 53 families
acting as pollen and nectar source for pollinators in the southern Brazil. They
reported that Asteraceae (15 spp.) was dominant family followed by Rubiaceae
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
11
(10 spp.), Fabaceae (7 spp.) and Solanaceae (6 spp.). Adhikari and Ranabhat
(2011) reported a total of 158 species as pollen and nectar producing plants for
pollinators from the mid-hills of central Nepal, out of the which, 19 species
were horticultural plants, 42 species were crop plants, 15 species were
ornamental plants and 82 species were wild plants. They further recognized 38
species as major, 35 as medium and 30 as minor sources for both nectar and
pollen. Bista (2001) found 119 plant species acting as foraging sites for
pollinators, out of which 47 species were found major sources for honeybees at
Kabre area of Dolakha district, Nepal. Ayansola and Davies (2012) found 49
species from the tropical rain forest and Savanna zones of south-western
Nigeria. Nnamani and Uguru (2012) found 56 species belonging to 53 genera
and 21 families, and reported that Fabaceae, Euphorbiaceae and Anacardiaceae
were the predominantly visited families by insect visitors in southeastern
Nigeria. Dukku (2013) found 61 plants acting as food sites for honey bees from
Savanna zone of north-eastern Nigeria. Wubie et al. (2014) found that 12 plant
species were found major source of pollen for pollinators as well as for bee
keeping in Ethiopia. They also found that majority of flora visited by
pollinators was herbaceous in nature.
Suryanarayana et al. (1992) found a total of 40 and 48 plant species as
foraging sites for Apis cerana and A. mellifera respectively from Muzaffarpur
district of Bihar, India. Datta et al. (2008) studied bee flora of the upper
Gangetic region of India and found that, in addition to already known bee flora,
Brassica, Coriandrum, Litchi and Eucalyptus and Callistemon which are
planted for social forestry programs are also utilized as pollen and nectar
sources by honey bees. Shubharani et al. (2012) found 91 species belonging to
42 flowering plant families as pollen and nectar sources for pollinators from
Coorg district of Karnataka, India. Recently, Bhalchandra et al. (2014)
recorded 52 plant species, including 29 agricultural crops and 23 wild plants,
from the biodiversity hotpot region of Western Ghats, India, that were useful to
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
12
honey bees. Gaur et al. (2014) recorded 521 bee forage plant species belonging
to 377 genera and 117 families from the Garhwal Himalaya, India. Behera et
al. (2014) found that Apis cerana and A. mellifera visited 25 plant species for
nectar and pollen.
The Himalayas represent one of the world’s tallest ranges of mountains,
which are located in the Indian subcontinent. Being one of the global
biodiversity hotspots, the Himalaya possess a precious biological wealth of
great economic value and scientific interest. Kashmir Himalaya, lying in the
north-western part of Himalyas, is renowned for its natural landscape
characterized by a diverse and endemic biodiversity (Lawrence, 1895; Dar et
al., 2013). Notwithstanding rich biodiversity, very few research studies have
been carried out to explore the diversity of flowering plants and their insect
visitors in this Himalayan region. Williams (1991) mentioned the presence of
29 species of flower visitors (bumble bees) from Kashmir Himalaya. Saini et
al. (2012) reported 27 species of flower visitors (bumble bees), belonging to 10
subgenera of the genus Bombus from Kashmir. Ganie et al. (2013) reported 17
species of flower visitors belonging to 11 families and 3 orders: Hymenoptera,
Diptera and Lepidoptera from the apple (Malus domestica) orchards of
Kashmir valley. Recently, Paray et al. (2014) recorded 15 species of flower
visitors belonging to 8 genera and 4 families and their host plants from the
apple orchards of Kashmir valley.
The preceding studies have been either narrow in scope, focussed on
insect visitors only without documenting the diversity of flowering plants in
detail or specific to cultivated plants such as apple. It is in this context that the
present study, using an integrated approach, attempts to document the diversity
of flowering plants and their insect visitors at a much broader scale spanning
across the wild and cultivated plants in the Kashmir Himalaya.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
13
STUDY AREA
Kashmir Himalaya represents a biotic province in the bio-geographic
zone of the North-Western Himalaya in India (Rodgers and Panwar 1988). It
lies between 33° 20′ to 34° 54′ N latitudes and 73° 55′ to 75° 35′ E longitudes.
The region mainly comprises of the famous valley of Kashmir, surrounded by
high mountain ranges on all sides (Husain, 2001). The climate is predominantly
temperate with wet and cold winters and relatively dry and moderately warm
summers. It is marked by well-defined seasonality, with four seasons a year,
winter (December-February), spring (March-May), summer (June-August) and
autumn (September-November). Different seasons have beautified Kashmir
with different flowers interacting with different insect visitors shaping different
flower-visitor architectural patterning with wide or narrow webs.
Figure 1.1: Location map of the study area
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
14
Materials and Methods
During the present study, systematic surveys and field observations were
carried out in different months of the year 2014-2015 in the study area (Fig.
1.1) to record data regarding pollen and nectar producing plants and flower
visitors visiting them. Geo-coordinates of each site were recorded by using
GPS (Garmin Montana 650). During each survey, field observations were
recorded at different sites in the study area. At each site, visual observations
were keenly recorded on the basis of visitor centred approach near the flowers
of a particular plant. Visitor centred approach was recorded on full bloomed
flowers, and the number of different flower visitors visiting the flower was
quantified (Herrera, 1988; Mahy et al., 1998; Dicks et al., 2002). The event of
flower-visitor interaction was photographed (Make: Sony, Model: DSC-S2000)
for further processing and study in the laboratory.
Plant specimens were collected for their correct scientific identification.
Using standard herbarium methodology (Bridson and Forman, 1998), plant
specimens were processed and identified on the basis of relevant taxonomic
literature (Stewart, 1972) and further validated with expert determination and
using specialized online websites (Flora of Pakistan @efloras.org; Flora of
China @efloras.org; www.flowersofindia.net).
The flower insect visitors were collected by collecting nets and were
killed in ethyl acetate solution, then stretched and pinned for identification
purposes. Labels were appended to each sample containing date of collection,
name of locality, altitude, longitude, latitude and collector’s name. The labelled
samples were placed in the boxes specially designed for the above-mentioned
purpose. Flower visitor specimens were identified on the basis of relevant
taxonomic literature (Michener 2007) and further validated with expert
determination by Kumar Ghorpede, Department of Entomology, Dharwad
Agricultural University, Hubli, Karnataka, India and Alain Pauly, Department
of Entomology, Royal Belgian Institute, Belgium (Photoplate 1.1).
The type of foraging plant was recognised on the basis of foraging
behaviour of flower visitor on the flower. These include: (i) when the flower
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
15
visitor stays at a particular position on a flower especially at the base of
petal/tepal and pierces its proboscis towards the position of nectary to suck the
nectar, then the plant was categorised as the nectar producing plant; (ii) when
flower visitor roams around the flower display near the anthers in different
patterns, especially in a ring pattern in bowl-shaped flowers, and collects pollen
on its body, then the plant was categorised as pollen producing plant; and (iii)
when both the preceeding foraging behaviours of insect visitor was observed
for the flowers of a particular plant, then the plant was categorised as both
pollen and nectar producing plant.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
16
RESULTS
Diversity of pollen and nectar producing plants
During the present study, a total of 227 flowering plant species
belonging to 182 genera and 58 families were recorded that are potential
foraging plants. Out of the total plant species, 180 herbs, 26 shrubs and 21 trees
were found to be acting as pollen and nectar producing plants for flower
visitors in the study area (Table 1.1 and Photoplates 1.2-1.5). Among the 58
plant families, Asteraceae contributed the largest number of 35 plant species,
followed by Rosaceae and Lamiaceae with 24 and 15 plant species
respectively. On the other hand there were 28 families which contributed only
single plant species each (Fig. 1.2). Majority of these foraging plants were
annual herbs (97 species), followed by 74 perennial herbs, 27 shrubs, 22 trees
and 7 biennial herbs (Fig. 1.3). Nearly two-third (i.e. 152 species) of the
recorded foraging plants investigated were wild growing species, and the
remaining 75 plant species were cultivated for different purposes: floriculture
(37 species), agriculture (19 species), horticulture (15 species) and social
forestry (04 species) (Fig. 1.4). During the present study, the foraging plants
were found to provide three different types of rewards for the insect visitors:
180 plant species provide both nectar and pollen as reward, 42 and 5 plant
species exclusively provide pollen and nectar to the flower visitors,
respectively (Fig. 1.5).
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
17
Diversity of flower visitors
The present study has recorded diverse insects that visit the flowers of
these foraging plants to obtain their food. A total of 70 flower visitors
belonging to 43 genera and 23 families were found visiting for their food to
above memtioned flowers (Table 1.2; Photoplates 1.6 and 1.7). The family
Apidae (12 species) was the dominant group of flower visitors followed by
Halictidae (11 species), Syrphidae (10 species), Andrenidae, Nymphalidae,
Megachalidae, Pieridae (04 species each), Ceratidae (03 species), Bibionidae,
Muscidae, Vespidae, Scathophagidae (02 species each) and rest of families:
Formacedae, Pyrrhocoridae, Calliphoridae, Hesperiidae, Lycaenidae,
Tenthredinidae, Sphingidae, Meloidae, Scarabaeidae, Coccinellidae, Thripidae
(01 species each) (Fig. 1.6)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
18
Table 1.1: Diversity of foraging plants, their family, English name, Kashmiri name (if avialbale), habit, purpose of cultivation, insect visitor
(s), reward, geo-coordinates and site of record in Kashmir Himalaya (*Abbreviation used: PN = Pollen and nectar producing
plants, PL = Pollen producing plants, NC = Nectar producing plants, # taxonomic names at the level of genus have been written for
insect visitors).
Species name Family English name
Kashmiri
name Habit Purpose Insect visitors
(spp.)# *Reward
Latitude
N
Longitude
E
Altitude
(m) *Recording Site
Abelia grandiflora
Rehd.
Caprifoliaceae Glossy Abelia Shrub Ornamental Apis, Bombus,
Xylocopa
PN 3408'45.07" 7452'36.83" 1606.46 SKUAST-K (SC)
Abelmoschus
esculentus Moench
Malvaceae Ladies
Finger
Annual herb Olericulture Apis, Bombus PN 3408'50.27" 7452'40.33" 1609.23 SKUAST-K (SC)
Aconitum laeve
Royle
Ranunculaceae Grape-leaved
Monkshood
Annual herb Wild Apis, Bombus PN 3419'23.41" 7444'07.33" 3195.38 Rangnar
Aconogonum molle
Hara
Polygonaceae Sikkim
Knotweed
Annual herb Wild Apis PL 3419'35.73" 7444'11.19" 3293.54 Gager Patri
Actinidia deliciosa
Liang and Ferguson.
Actinidiaceae Kiwi Kiwi Climber Horticulture/
Ornamental
Apis PN 3408'47.65" 7452'55.70" 1621.54 SKUAST-K (SC)
Aesculus indica
Hook.
Sapindaceae Indian horse
Chestnut
Tree Wild Apis PN 3419'58.05" 7441'09.03" 1703.08 SKUAST-K (SC)
Ageratum
houstonianum Mill.
Asteraceae Floss Flower Annual herb Ornamental Apis, Bombus PN 3408'49.29" 7452'58.37" 1623.08 SKUAST-K (SC)
Agrimonia pilosa
Ledeb.
Rosaceae Hairy Agrimony Perennial
herb
Wild,
Medicinal
Apis PN 3419'06.68" 7444'11.55" 3276 Gager Patri
Ailanthus altissima
Swingle
Scrophulariaceae Tree of Heaven Allum
Thrass
Tree Social
Forestry
Apis PN 3408'62.15" 7453'26.39" 1639.69 Gager Patri
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
19
Albizia julibrissin
Durazz.
Fabaceae Silk tree mimosa Tree Landscape Apis PL 3408'48.28" 7452'37.42" 1606.46 SKUAST-K (SC)
Alcea rosea L. Malvaceae Hollyhock Perennial
herb
Ornamental Apis, Bombus PN 3408'49.32" 7452'58.19" 1626.15 SKUAST-K (SC)
Allium cepa L. Alliaceae Onion Gande Biennial herb
Agriculture Apis, Ceratina
Lasioglossum,
Halictus
PN 3408'56.26" 7452'49.41" 1614.78 SKUAST-K (SC)
Allium sativum L. Alliaceae Garlic Rohon Biennial herb
Agriculture Apis, Ceratina
PN 3408'55.82" 7452'48.88" 1614.78 SKUAST-K (SC)
Anaphalis busua
DC.,
Asteraceae Perennial
herb
Wild Apis PN 3419'01.52" 7443'43.34" 3357.23 Baj patri
Anemone obtusiloba
Don
Ranunculaceae Himalayan
thimble weed
Annual herb Wild Apis PN 3419'07.33" 7443'57.27" 3127.69 Gager Patri
Anemone tetrasepala
Royle
Ranunculaceae Four Petal
anemone
Annual herb Wild Apis PN 3419'12.39" 7443'53.83" 3081.54 Gager Patri
Anthemis cotula L. Asteraceae Sticking
Chamomile
Faggass Annual herb Wild Apis,
Metasyrphus,
Sphaerophori,
Episyrphus,
Eristalodes,
Eristalis,
Eoseristalis,
Thrips.
PN 3408'50.72" 745301.79" 1629.54 SKUAST-K (SC)
Antirrhinum majus
L.
Scrophulariaceae Dog Flower Annual herb Ornamental
Xylocopa. Apis,
Ceratina
PN 3408'48.14" 7452'55.91" 1623.08 SKUAST-K (SC)
Aquilegia fragrans Ranunculaceae Fragrant Perennial Wild Apis PN 3419'16.52" 7444'08.56" 3234.46 Baj patri
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
20
Benth. Columbine herb
Arabis glabra Bernh.
Brassicaceae Tower Mustard Perennial
herb
Wild Apis,
Lasioglossum
PN 3338'30.44" 7446'48.20" 2308.62 Ahrabal
Arctium lappa L. Asteraceae Greater Burdock Perennial
herb
Wild Xylocopa,
Ceratina
PL 3408'45.26" 7452'00.84" 1599.7 SKUAST-K (SC)
Artemisia
absinthium L.
Asteraceae Wormwood Annual herb Wild,
Medicinal
Apis PL 3408'49.52" 7453'02.21" 1635.69 SKUAST-K (SC)
Aster thomsonii
Clarke
Asteraceae Blue aster Annual herb Lasioglossum PN 3419'09.44" 7444'15.17" 3357.23 Gager Patri
Astragalus
grahamianus Benth.
Fabaceae Milk- Vetch Perennial
herb
Wild Amegilla,
Andrena,
Formica
PN 3410'29.81" 7454'34.27" 1828.62 Dara
Bellis perennis L. Asteraceae Perennial
herb
Wild Apis, Thrips PN 3408'49.42" 7453'02.30" 1636.31 SKUAST-K (SC)
Berberis lycium
Royle
Berberidaceae Indian barberry Kaw
Daach
Shrub Wild,
Medicinal
Apis PN 3419'51.52" 7441'01.73" 1667.69 SKUAST-K (SC)
Bergenia ligulata
Engl.
Saxifragaceae Winter Begonia Palpapper
Perennial
herb
Wild Apis, Bombus PN 3418'43.84" 7444'07.99" 3398.15 Gager Patri
Bidens tripartita L. Asteraceae Marigold bur Annual herb Wild Sphaerophoria PL 3408'46.57" 7452'46.72" 1611.4 SKUAST-K (SC)
Brassica compestris
L.
Brassicaceae Mustard Tilgogul Biennial herb
Olericulture Apis,
Scathophagia
PN 3408'54.07" 7452'50.78" 1616.62 SKUAST-K (SC)
Brassica rapa L. Brassicaceae Raddish Biennial herb
Olericulture Apis PN 3408'54.70" 7452'51.91" 1616.62 SKUAST-K (SC)
Buddleja davidii
Franch.
Scrophulariaceae Butterfly brush Shrub Ornamental Apis, Pieris,
Vanessa
PN 3408'52.87" 7452'37.05" 1607.38 SKUAST-K (SC)
Calendula officinalis Asteraceae Common Annual herb Ornamental Apis PN 3408'48.69" 7452'56.96" 1623.08 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
21
L. Marigold
Caltha alba
Cambess.
Ranunculaceae White Marsh
Marigold
Perennial
herb
Wild Apis,
Lasioglossum
PN 3419'09.16" 7443'53.58" 3127.69 Gager Patri
Campanula
cashmeriana Royle
Campanulaceae Kashmir
Bellflower
perennial
herb
Wild Apis,
Lasioglossum
PN 3419'16.56" 7443'43.09 3046.77 Rangnar
Campanula
rotundifolia L.
Campanulaceae Alpine bluebell Perennial
herb
Wild Apis,
Lasioglossum
PN 3419'17.55" 7433'33.19 3077.85 Rangnar
Campsis grandiflora
Schum.
Bignonaceae Trumpet Shrub Ornamental Apis, Bombus,
Xylocopa
PN 3408'48.43" 7452'28.27" 1604 SKUAST-K (SC)
Capsella bursa-
pastoris Medik.
Brassicaceae Shepherd's Purse Annual herb Wild Sphaerophoria PN 3408'51.48" 7453'00.01" 1622.46 SKUAST-K (SC)
Carduus edelbergii
Rech.f.
Asteraceae Thistle Perennial
herb
Wild Apis, Bombus,
Xylocopa
PN 3408'56.12" 7452'56.30" 1620.62 SKUAST-K (SC)
Carya illinoinensis
Koch
Juglandaceae Pecan nut tree Tree Horticulture Apis,
Scathophagia
PL 3408'48.40" 7453'01.30" 1636.62 SKUAST-K (SC)
Castanea sativa
Mill.
Fagaceae Chestnut Tree Horticulture Apis PL 3408'47.09" 7452'54.76" 1619.69 SKUAST-K (SC)
Centaurea cyanus L.
Asteraceae Corn flower Annual herb Ornamental Apis PL 3408'47.96 7452'56.54" 1623.08 SKUAST-K (SC)
Centaurea iberica
Spreng.
Asteraceae Iberian star
Thistle
Annual herb Wild Apis PN 3420'19.98" 7441'59.81" 1984.92 Ajas Bandipore
Cercis canadensis L.
Fabaceae Eastern Redbud Tree Wild Apis, Bombus,
Xylocopa
PN 3408'47.22" 7452'42.46" 1606.15 SKUAST-K (SC)
Chenopodium album
L.
Amaranthaceae Bathua Annual herb Wild Apis PL 3408'47.93" 7452'57.80" 1626.77 SKUAST-K (SC)
Chrysanthemum Asteraceae Crown Daisy Annual herb Ornamental Apis, PN 3408'48.20" 7452'57.23" 1623.08 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
22
coronarium Spach Coccinella
Cichorium intybus L.
Asteraceae Chicory Perennial
herb
Wild Apis, Bombus,
Sphaerophoria
PN 3408'48.70" 7452'51.07" 1614.15 SKUAST-K (SC)
Cirsium arvense
Scop.
Asteraceae Wool bearing
Thistle
Perennial
herb
Wild Apis, Bombus,
Xylocopa
PN 3408'56.06" 7452'56.56" 1620.62 SKUAST-K (SC)
Cirsium falconeri
Petr.
Asteraceae Falconer's Thistle
Perennial
herb
Wild Apis, Bombus
Xylocopa
PN 3408'43.63" 7452'43.63" 1637.85 SKUAST-K (SC)
Cirsium vulgare
Ten.
Asteraceae Thistle Perennial
herb
Wild Apis, Bombus,
Xylocopa
PN 3420'11.60" 7441'36.64" 1848.62 Ajas Bandipore
Clematis montana
DC.
Ranunculaceae Anemone
clematis
Annual herb Wild Apis PN 3419'39.44" 7443'23.23" 2729.23 Rangnar
Codonopsis ovata
Benth.
Campanulaceae Kashmir Bonnet
Bellflower
Perennial
herb
Wild Apis,
Lasioglossum
PN 3419'25.56" 7443'43.23" 3044.92 Rangnar
Colchicum luteum L.
Colchicaceae Yellow
Colchicum
Perennial
herb
Wild Apis,
Scathophagia
PN 3408'49.24" 7453'02.68" 1638.77 SKUAST-K (SC)
Conium maculatum
L.
Apiaceae Poison hemlock Gadn
Baidion
Perennial
herb
Wild Sphaerophoria,
Calliphora
PL 3409'00.99" 7453'00.03" 1624 SKUAST-K (SC)
Convolvulus arvense
L.
Convolvulaceae Field Bindweed Perennial
herb
Wild Apis PN 3408'50.98" 745302.04" 1629.54 SKUAST-K (SC)
Coriandrum sativum
L.
Apiaceae Coriander Dhaniwal
Annual herb Olericulture Apis, Andrena PL 3408'50.59" 7452'43.18" 1608.61 SKUAST-K (SC)
Corydalis
cashmeriana Royle
Papaveraceae Kashmir
Corydalis
Perennial
herb
Wild Apis PN 3419'05.04" 744339.34" 3240.62 Gager Patri
Corydalis diphylla
Wall.
Papaveraceae Two leaved
corydalis
Annual herb Wild Apis PN 3419'05.04" 7443'39.34" 3240.62 Rangnar
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
23
Crepis tectorum L. Asteraceae Narrow-leaved
Hawk's-beard
Annual herb Wild Apis PL 3408'58.15" 7452'57.95" 52721622 SKUAST-K (SC)
Cucumis melo L. Cucurbitaceae Wild Melon Annual herb Olericulture Apis PN 3408'51.79" 7452'46.99" 1611.38 SKUAST-K (SC)
Cucumis sativus L. Cucurbitaceae Cucumber Annual herb Olericulture Apis PN 3408'51.84" 7452'46.96" 1611.38 SKUAST-K (SC)
Cucurbita maxima
Duchesne
Cucurbitaceae Giant pumpkin Annual herb Olericulture Apis PN 3408'50.07" 7453'02.30" 1633.54 SKUAST-K (SC)
Cucurbita pepo L. Cucurbitaceae Pumpkin Annual herb Olericulture Apis PN 3408'49.99" 7453'02.39" 1633.54 SKUAST-K (SC)
Cydonia oblonga
Miller
Rosaceae Quince Bomb
Choont
Tree Horticulture Apis,
Lasioglossum
PN 3408'47.15" 7452'43.33" 1607.38 SKUAST-K (SC)
Cynodon dactylon
Pers.
Poaceae Bermuda grass Dramun Perennial
herb
Wild Apis PL 3408'48.16" 7452'41.18" 1606.46 SKUAST-K (SC)
Cynoglossum
glochidiatus Wall
Boraginaceae Annual herb Wild Apis PN 3408'53.16" 7452'59.10" 1618.46 Longmarg
Datura stramonium
L.
Solanaceae Jimsonweed Annual herb Wild Episyrphus
balteatus
PN 7452'33.20" 7452'42.09" 1597.85 SKUAST-K (SC)
Daucus carota L. Apiaceae Carrot Gazer Biennial herb
Horticulture Apis, Formica,
Musca
PL 3408'58.77" 7452'57.94" 1622.15 SKUAST-K (SC)
Delphinium roylei
Munz
Ranunculaceae Royle's Larkspur Perennial
herb
Ornamental Apis PN 3419'38.52" 7443'27.87" 2779.38 Baj Patri
Descurainia sophia
Prantl
Brassicaceae Herb sophia/ Fix
weed
Annual herb Wild Apis PN 3408'50.06" 7452'58.43" 1623.38 SKUAST-K (SC)
Digitalis purpurea
L.
Scrophulariaceae Foxgloves Perennial
herb
Ornamental Apis PN 3408'47.75" 7452'56.45" 1623.08 SKUAST-K (SC)
Dipsacus inermis
Wall.
Dipsacaceae Himalayan teasel
Perennial
herb
Wild Bombus PN 3419'38.19" 7443'24.42" 2740 Mirnar
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
24
Doronicum falconeri
Hook.
Asteraceae leopard's bane Perennial
herb
Wild Lasioglossum PN 3419'08.62" 7443'35.09" 3201.85 Rangnar
Echium
plantagineum L.
Boraginaceae Purple Viper's
Bugloss
Annual herb Ornamental Apis PN 3408'45.74" 7452'36.66" 1606.77 SKUAST-K (SC)
Epilobium hirsutum
L.
Onagraceae Hairy Willow
herb
Annual herb Wild Apis PN 3408'46.46" 7452'41.59" 1606.46 SKUAST-K (SC)
Epilobium
parviflorum Schreb.
Onagraceae Small flowered
willow herb
Annual herb Wild Apis PN 3420'00.59" 7443'42.50" 3142.46 Ajas Bandipore
Epilobium
royleanum Hausskn.
Onagraceae Royle's Willow
herb
Annual herb Wild Apis PN 3408'56.58" 7452'56.06" 1620.31 SKUAST-K (SC)
Eremurus
himalaicus Baker.
Liliaceae Himalayan
Desert Candle
Perennial
herb
Wild Apis PN 3420'22.58" 7442'07.18" 2049.54 Ajas Bandipore
Eriobotyria japonica
Lindl.
Rosaceae Loquat Loquat Tree Horticulture Apis, Bombus.
Xylocopa,
Lasioglossum
.
PN 3408'47.37" 7452'42.12 1606.15 SKUAST-K (SC)
Erysimum
hieraciifolium L.
Brassicaceae European
wallflower
Perennial
herb
Wild Apis,
Lasioglossum
.
PN 3408'59.59" 7451'01.47 1624 SKUAST-K (SC)
Eschscholzia
californica Cham.
Papaveraceae California Poppy
Perennial
herb
Ornamental Apis PN 3408'47.65" 7452'56.35" 1623.08 SKUAST-K (SC)
Euonymus
hamiltonianus Wall.
Celastraceae Himalayan
Spidle tree
Perennial
shrub
Wild Apis PN 3408'44.98" 7452'41.75" 1606.46 SKUAST-K (SC)
Euphorbia
helioscopia L.
Euphorbiaceae Sun Spurge Zuri
Sochel
Annual herb Wild Apis,
Sphaerophoria
PN 3408'49.88" 745300.81" 1629.23 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
25
Euphorbia wallichii
Hook.f.
Euphorbiaceae Wallich Spurge Annual herb Wild Apis,
Scathophagia
PN 341937.57" 744346.16" 3004.62 Longmarg
Foeniculum vulgare
Mill.
Apiaceae Fennel Baidion Perennial
herb
Olericulture Waspa,
Formica
PN 3408'46.50" 7452'41.18" 1606.15 SKUAST-K (SC)
Forsythia
viridissima Lindl.
Oleaceae Golden bells Shrub Ornamental Apis,
Scathophagia
PN 3408'46.05" 7452'37.42" 1607.08 SKUAST-K (SC)
Fragaria ananassa
Duchesne
Rosaceae Strawberry Strawberr
y
Perennial
herb
Horticulture Apis,
Lasioglossum
PN 3408'53.42" 7452'57.07" 1622.15 SKUAST-K (SC)
Fragaria nubicola
Lacaita
Rosaceae Himalayan
strawberry
Perennial
herb
Wild Apis,
Lasioglossum.
PN 3339'20.25" 7446'29.38" 2463.69 Ahrabal
Fumaria indica
Pugsley
Papaveraceae Indian Fumitory Annual herb Wild,
Medicinal
Apis NC 3408'51.33" 7452'56.10" 1617.85 SKUAST-K (SC)
Galinsoga parviflora
Cav.
Asteraceae Quick weed Annual herb Wild Sphaerophoria PL 3408'50.44" 7452'56.95" 1620 SKUAST-K (SC)
Gentiana
cachemirica Decne.
Gentianaceae Dwarf Willow
Gentian
Perennial
herb
Wild Bombus PN 3418'32.22" 7443'52.91" 3376.92 Maal
Geranium nepalense
Sweet
Geraniaceae Annual herb Wild Apis PN 3408'52.61" 7452'58.65" 1621.23 SKUAST-K (SC)
Geranium pratense
L
Geraniaceae Meadow
Geranium
Annual herb Wild Apis PN 3419'30.87" 7443'12.97" 2829.85 Mirnar
Geranium
wallichianum Don
Geraniaceae Wallich
Geranium
Annual herb Wild Apis PN 3418'33.66" 7443'09.11" 2798.77 Shal Patri
Geum roylei Wall. Rosaceae Royle's Avens Annual herb Wild Apis, Waspa PN 3408'57.11" 7452'57.13" 1621.54 SKUAST-K (SC)
Gladiolus hortulanus
Bailey
Iridaceae Garden Gladiola/
Sword lily
Annual herb Ornamental Apis PN 3408'49.11" 7452'58.63" 1623.08 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
26
Hackelia uncinata
Fisch.
Boraginaceae Hooked stickseed
Perennial
herb
Wild Apis. PN 3419'33.70" 744324.00" 2782.46 Baj Patri
Halianthus annus L. Asteraceae Common
sunflower
Guliaftab Annual herb Ornamental Apis, Xylocopa
PN 3408'48.85" 7452'57.11" 1622.77 SKUAST-K (SC)
Heracleum
candicans Wall.
Apiaceae White leaf
Hogweed
Biennial herb
Wild Apis, Formica PL 3419'31.25" 7443'10.86" 2830.15 Gager Patri
Hibiscus rosa-
sinensis L.
Malvaceae China rose Shrub Ornamental Apis, Bombus,
Xylocopa
PN 3408'46.40" 7452'43.40" 1607.08 SKUAST-K (SC)
Hypericum
hookerianum Wight
and Arn.
Hypericaceae Johnswort Shrub Ornamental Apis, Bombus
Xylocopa
Lasioglossum
PN 3408'46.59" 7452'41.28" 1606.46 SKUAST-K (SC)
Hypericum
perforatum L.
Hypericaceae Perforate Johns
Wort
Annual herb Wild Apis PN 3408'59.54" 7452'59.97" 1623.38 SKUAST-K (SC)
Iberis amara L. Brassicaceae Rocket Candytuft
Annual herb Ornamental Apis PN 3408'48.10" 7452'57.19" 1624.31 SKUAST-K (SC)
Impatiens
glandulifera Royle
Balsaminaceae Himalayan
balsam
Annual herb Wild Bombus PN 3419'58.00" 7441'02.10" 1684 Ajas Bandipore
Indigofera
heterantha Wall
Fabaceae Himalayan
indigo
Shrub Wild Apis PN 3419'56.08" 7442'37.80" 2496.31 Baj patri
Inula royleana Clark
Asteraceae Annual herb Wild Apis PN 3418'41.60" 7444'48.22" 3518.77 Gager Patri
Inula racemosa
Hook.
Asteraceae Pushkarmool Elecampa
ne
Annual herb Wild Apis PN 3408'44.52" 7452'54.26" 1622.15 SKUAST-K (SC)
Ipomoea tricolor
Cav.
Convolvulaceae Common glory Perennial
herb
Wild Apis PN 3408'48.44" 7452'54.61" 1617.54 SKUAST-K (SC)
Iris decora Wall. Iridaceae Graceful
Himalayan Iris
Mazzar
Mund
Perennial
herb
Ornamental Xylocopa,
Lasioglossum
PN 3408'49.12" 7452'58.09" 1624.62 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
27
Iris hookeriana
Foster
Iridaceae Hooker's Iris Mazzar
Mund
Perennial
herb
Ornamental Xylocopa,
Lasioglossum
PN 3408'45.09" 7452'57.65" 1635.08 SKUAST-K (SC)
Lactuca
dolichophylla Kitam.
Asteraceae Long-leaved
lettuce
Annual herb Wild Apis,
Sphaerophoria
PL 3408'58.61" 7452'58.33" 1622.46 SKUAST-K (SC)
Lagotis cashmeriana
Rupr.
Scrophulariaceae Kashmir Lagotis
Perennial
herb
Wild Apis
Bombus
PN 3420'00.95" 7443'47849"
3183.38 Gager Patri
Lamium album L. Lamiaceae White dead nettle
Annual herb Wild Apis PN 3408'49.31" 7452'55.45" 1618.46 SKUAST-K (SC)
Lavandula officinalis
L.
Lamiaceae Lavender Perennial
herb
Ornamental Apis PL 3408'45.46" 7452'39.67" 1606.15 SKUAST-K (SC)
Lavatera
cashmiriana
Cambess.
Malvaceae Tree Mallow Sazeposh Perennial
herb
Ornamental Apis , Xylocopa
PN 3408'49.41" 7452'35.77" 1607.38 SKUAST-K (SC)
Leonurus cardiaca
L.
Lamiaceae Motherwort Perennial
herb
Wild Apis PN 3419'02.50" 744340.98" 3243.38 Baj Patri
Ligularia fischeri
Turcz.
Asteraceae leopard plant Perennial
herb
Wild Apis, Bombus PN 3418'39.38" 7444'21.80" 3451.38 Gager Patri
Lindelofia longiflora
Baill.
Boraginaceae Himalayan
lungwort
Perennial
herb
Wild Apis, Bombus
Xylocopa
PN 3420'12.67" 7443'55.45" 3419.15 Longmarg
Lonicera japonica
Thunb.
Caprifoliaceae Honey suckle Shrub Ornamental Apis PN 3408'56.96" 7453'04.39" 1625.54 SKUAST-K (SC)
Lotus corniculatus
L.
Fabaceae Common birds
foot trefoil
Annual herb Wild Apis PN 3408'58.73" 7453'00.97" 1623.38 SKUAST-K (SC)
Lycopersicon
esculentum Mill.
Solanaceae Tomato Annual herb Olericulture Apis, Bombus PN 3408'51.41" 7452'46..53"
1611.07 SKUAST-K (SC)
Magnolia Magnoliaceae Bull Tree Ornamental Apis, PN 3408'44.18" 7452'41.08" 1606.15 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
28
grandiflora L. Bay/Southern
Magnolia
Scathophagia
Mahonia borealis
Takeda
Berberidaceae Perennial
herb
Wild Apis PN 340751.03" 7450'01.54" 1606.46 Kashmir
University
Malus domestica
Borkh.
Rosaceae Apple Choont Tree Horticulture Apis, Xylocopa
Andrena.
Bombus,
PN 3408'50.67" 7452'48.36" 1612.31 SKUAST-K (SC)
Malva neglecta
Wallr.
Malvaceae Common Mallow
Sochel Annual herb Wild Apis PN 3408'51.03" 7452'01.98" 1628.92 SKUAST-K (SC)
Malva sylvestris L. Malvaceae High Mallow Sochel Annual herb Wild Apis PN 3408'49.96" 7452'58.52" 1623.69 SKUAST-K (SC)
Marrubium vulgare
L
Lamiaceae Horehound Perennial
herb
Wild Apis PN 3408'31.64" 7451'43.50" 1596 SKUAST-K (SC)
Meconopsis latifolia
Prain
Papaveraceae Blue Poppy Annual herb Wild,
Medicinal
Apis PN 3418'43.48" 7444'47.89" 11570356
0
Gager Patri
Medicago lupulina
L.
Fabaceae Black medic Annual herb Wild Apis PL 3408'41.83" 7452'71.11" 1613.85 SKUAST-K (SC)
Mentha arvensis L. Lamiaceae Field Mint Perennial
herb
Olereculture Apis PN 3408'58.18" 7452'53.56" 1616.92 SKUAST-K (SC)
Mentha longifolia L.
Lamiaceae Horse Mint Perennial
herb
Wild Apis, Waspa PN 3408'48.18" 7452'53.96 1617.23 SKUAST-K (SC)
Morina longifolia
DC.
Morinaceae Nepalese Whorl
flower
Perennial
herb
Wild Apis, Bombus PN 3419'02.04" 7443'42.45" 3253.85 Baj Patri
Myosotis arvensis L.
Boraginaceae Forget me not Annual herb Wild Apis PN 3339'21.39" 7446'32.17" 2448 Ahrabal
Myosotis caespitosa
Schultz
Boraginaceae Annual herb Wild Apis PN 3339'21.48" 7446'32.95" 2449.23 Ahrabal
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
29
Nepeta erecta Benth.
Lamiaceae Erect Catmint Annual herb Wild Apis,
Polyommatus
PN 3408'52.33" 7452'53.25" 1619.38 SKUAST-K (SC)
Narcissus poeticus
L.
Amaryllidaceae Daffodil Yember
Zal
Perennial
herb
Ornamental Apis,
Scathophagia
PN 3408'45.05" 7452'57.72" 1635.38 SKUAST-K (SC)
Nasturtium officinale
Aiton
Brassicaceae Watercress Perennial
herb
Wild Apis PN 3408'48.20" 7452'54.18" 1616.92 SKUAST-K (SC)
Nelumbo nucifera
Gaertn.
Nelumbonaceae Lotus Pamposh Perennial
herb
Wild,
Aquatic
Apis PN 3415'19.54" 744109.15" 1596 Mansbal Lake
Nerium indicum
Mill.
Apocynaceae Oleander Shrub Ornamental Apis PN 3408'45.02" 7452'38.05 1606.46 SKUAST-K (SC)
Nymphaea alba L. Nymphaeaceae Water Lily Perennial
herb
Wild,
Aquatic
Lasioglossum PN 3408'34.31" 7452'35.02" 1596 Dal Lake
Nymphoides peltata
Kuntze
Menyanthaceae Fringed water lily
Khur Perennial
herb
Wild,
Aquatic
Diptera PN 3408'34.28" 7452'38.04" 1596 Dal Lake
Oxalis corniculatus
L.
Oxalidaceae Sleeping beauty Chok
cheeni
Annual herb Wild Apis PN 3408'49.38" 7452'42.83" 1608 SKUAST-K (SC)
Oxytropis
cashmeriana
Cambess.
Fabaceae Locoweed Annual herb Wild Macroglossum
PN 3420'00.45" 7443'47.49" 3182.77 Gager Patri
Paparver dubium L. Papaveraceae Long headed
Poppy/ Blind
eyes
Guli lal Annual herb Wild Apis PN 3408'49.37" 7452'42.80" 1608 SKUAST-K (SC)
Papaver somniferum
L.
Papaveraceae Opium Poppy Annual herb Olereculture Apis PN 3408'48.60" 7452'57.90" 1623.08 SKUAST-K (SC)
Parthenium Asteraceae Carrot grass Perennial Wild Formica PL 3408'49.83" 7452'01.42" 1631.08 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
30
hysterophorus L. herb
Pedicularis
pyramidata Benth.
Orobanchaceae Pyramid
Lousewort
Perennial
herb
Wild Macroglossum
PN 3420'10.87" 7443'50.07" 3364 Longmarg
Petunia alba
Ferguson and Ottley
Solanaceae Petunia Annual herb Ornamental Pieris
brassicae L
PN 3408'45.46" 7452'36.24" 1606.77 SKUAST-K (SC)
Phaseolus vulgaris
L.
Fabaceae Beans Rajmah Annual herb Horticulture Apis, Bombus,
Xylocopa
PN 3408'52.60" 7452'47.75" 1611.38 SKUAST-K (SC)
Philadelphus
incanus Koehne
Hydrangeaceae Dogwood Shrub Wild Apis, Xylocopa
PN 34120'04.79"
74142'09.24"
2205.54 Mirnar
Phlomis bracteosa
Benth.
Lamiaceae Purple Jeruselum
Sage
Perennial
herb
Wild Apis PN 3419'01.87" 7443'41.86" 3367.07 Baj patri
Physalis
philadelphica Lam.
Solanaceae Ground cherry Annual herb Horticulture Apis, Bombus PN 3408'50.28" 7452'57.83" 1622.15 SKUAST-K (SC)
Pinus wallichiana A.
B. Jacks.
Pinaceae Himalayan Pine Tree Wild Bombus PL 3418'35.08" 7443'48.06" 3367.08 Malal
Plantago lanceolata
L.
Plantaginaceae Ribwort Plantain Annual herb Wild Apis PL 3408'47.07" 7452'46.99" 1611.69 SKUAST-K (SC)
Plantago major L. Plantaginaceae Broad leaf
Plantain
Annual herb Wild Apis PL 3408'49.76" 7453'01.25" 1631.08 SKUAST-K (SC)
Plectranthus rugosus
Benth.
Lamiaceae Wrinkled leaf
isodon
Sol Lie Shrub Wild Apis, Bombus PN 3420'17.12" 7442'22.88" 2142.77 Mirnar
Poa annua L. Poaceae Annual Bluegrass
Annual herb Wild Apis PL 3408'48.17" 7452'41.17" 1606.46 SKUAST-K (SC)
Podophylum
haxandrum Royle
Berberidaceae Himalayan May
Apple
Vanvagun
Perennial
herb
Wild,
Medicinal
Lasioglossum,
Halictus
PN 342003.11" 744451.06" 3075.69 Grat Watun
Polygonum Polygonaceae Water smartweed
Annual herb Wild Apis PL 3408'51.21" 7452'50.45" 1614.77 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
31
amphibium L.
Polygonum
amplexicaule Don
Polygonaceae Red Mountain
Fleece flower
Annual herb Wild Apis PL 3432'58.80" 7419'22.98" 1696.62 Kupwara
Potentilla
argyrophylla Lehm.
Rosaceae Silver leaved
cinquefoil
Perennial
herb
Wild Apis,
Lasioglossum
Scathophagia
PN 3340'11.21" 7502'04.30" 1703.69 Kulgam
Potentilla reptans L.
Rosaceae Creeping
cinquefoil
Perennial
herb
Wild Apis,
Lasioglossum
PN 3408'49.07" 7452'50.53" 1614.15 SKUAST-K (SC)
Prunella vulgaris L. Lamiaceae Common Self-
heal
Kalvut Perennial
herb
Wild Apis PN 3408'50.99" 7452'57.49" 1620.31 SKUAST-K (SC)
Prunus armeniaca L.
Rosaceae Apricot Cheere Tree Horticulture
Apis, Xylocopa,
Lasioglossum,
Scathophagia,
Ophyra
PN 3419'51.20" 744106.96" 1680.31 Ajas Bandipore
Prunus avium L. Rosaceae Cherry Gelas Tree Horticulture Apis, Xylocopa,
Scathophagia,
Lasioglossum,
Ophyra
PN 3408'55.71" 7452'57..65"
1621.85 SKUAST-K (SC)
Prunus domestica L.
Rosaceae Plum Aar Tree Horticulture Apis,
Xylocopa,
Scathophagia,
Lasioglossum,
Bibio, Ophyra
PN 3408'50.67" 745259.96" 1626.15 SKUAST-K (SC)
Prunus persica
Batsch
Rosaceae Peach Chunun Tree Horticulture Apis,
Xylocopa,
PN 3408'55.03" 7453'00.41" 1621.85 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
32
Lasioglossum ,
Ophyra
Pseudomartensia
nemorosa Stewart
and Kazmi
Boraginaceae Forest Alpine
Bluebell
Perennial
herb
Wild Bombus,
Macroglossum,
Mylabris
PN 34V19'25.32"
7443'40.73" 2919.08 Rangnar
Pteracanthus
urticifolius Bremek.
Acanthaceae Blue nettle Perennial
herb
Wild Apis PN 3408'52.66" 7452'59.74" 1621.54 SKUAST-K (SC)
Punica granatum L. Lythraceae Pome granate Tree Horticulture Apis PN 3408'50.03" 745300.04 1629.54 SKUAST-K (SC)
Pyrus communis L. Rosaceae Pear Tang Tree Horticulture Apis,
Xylocopa,
Lasioglossum,
Bibio, Plecia
PN 3408'49.29" 7452'59.20" 1626.77 SKUAST-K (SC)
Pyrus pyrifolia
Nakai
Rosaceae Pear Tang Tree Horticulture
Apis, Xylocopa,
Lasioglossum,
Bibio, Plecia,
Scathophagia
PN 3408'49.48" 7452'51.04" 1614.46 SKUAST-K (SC)
Ranunculus arvensis
L.
Ranunculaceae Corn Buttercup Annual herb Wild Apis PN 3409'02.72" 7453'01.90" 1624.92 SKUAST-K (SC)
Ranunculus laetus
Wall.
Ranunculaceae Cheerful
Buttercup
Perennial
herb
Wild Apis PN 3409'03.02"' 7453'03.56" 1625.23 SKUAST-K (SC)
Ranunculus
scleratus L.
Ranunculaceae Cursed Buttercup
Annual herb Wild Apis PN 3408'51.27" 7452'50.39" 1614.77 SKUAST-K (SC)
Raphanus sativus L.
Brassicaceae Turnip Biennial herb
Olericulture Apis PN 3408'54.13" 745250.83 1616.62 SKUAST-K (SC)
Rhododendron
campanulatum D.
Ericaceae Bell
Rhododendron
Shrub Wild Bombus PN 381847.50" 744353.39" 3285.54 Gager Patri
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
33
Don
Robinia
pseudoacacia L
Fabaceae Black Locust kiker Tree Social
forestry
Apis PN 3408'59.21" 7453'08.17" 1633.54 SKUAST-K (SC)
Rorippa islandica
Borbas
Brassicaceae Marsh yellow
cress
Annual herb Wild Apis PN 3408'50.35" 7452'58.07" 1622.46 SKUAST-K (SC)
Rosa brunonii Lindl.
Rosaceae Himalayan Musk
Rose
Shrub Wild Apis, Bombus
Xylocopa,
Lasioglossum
PN 3408'56.81" 7453'04.45" 1625.85 SKUAST-K (SC)
Rosa canina L. Rosaceae Dog Rose Shrub Wild Apis, Bombus,
Xylocopa,
Lasioglossum,
Cetonia
PN 3419'51.21" 7441'01.01" 1665.23 Ajas Bandipore
Rosa indica L. Rosaceae Rose Shrub Wild Apis, Bombus,
Xylocopa,
Cetonia
PN 3408'48.66" 7452'56.18" 1620.62 SKUAST-K (SC)
Rubus elipticus Sm. Rosaceae Yellow
Himalayan
Raspberry
Shrub Wild Apis, Ceratina,
Coccinella,
PN 3408'49.49" 7453'02.01" 1634.77 SKUAST-K (SC)
Rubus fruticosus L. Rosaceae Blackberry Shrub Wild Apis, Ceratina
PN 3408'51.28" 7452'02.26" 1628.92 SKUAST-K (SC)
Rubus niveus Thunb.
Rosaceae Raspberry Ganch Shrub Wild Apis, Ceratina
hieroglyphica
Smith
PN 3420'01.34" 7441'08.24" 1708.31 Ajas Bandipore
Rubus occidentalis
L.
Rosaceae Shrub Wild Apis, Ceratina PN 3408'51.04" 7452'58.98" 1622.46 SKUAST-K (SC)
Rubus ulmifolius Rosaceae Elm leaf Shrub Wild Apis, Ceratina
PN 3408'51.26" 7453'02.11" 1628.61 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
34
Schott blackberry
Rudbeckia hirta L. Asteraceae Black eyed Susan
Annual herb. Ornamental Apis, Bombus PN 3408'48.51" 7452'56.46" 1621.85 SKUAST-K (SC)
Rumex acetosa L. Polygonaceae Common Sorrel Abeeg Annual herb Wild Apis PL 3408'59.63" 7452'58.99" 1623.08 SKUAST-K (SC)
Salix alba L. Salicaceae Willow Veer Tree Social
Forestry
Apis,
Episyrphus
PL 3408'55.88" 7452'10.65" 1662.15 SKUAST-K (SC)
Salix caprea L. Salicaceae Goat willow Tree Landscape Apis PL 3407'53.45" 7450'03.89" 1605.23 Ajas Bandipore
Salvia hians Benth. Lamiaceae Himalayan Blue
Sage
Perennial
herb
Wild Apis, Bombus
Macroglossum
PN 3408'50.95" 7452'55.37" 1616.92 SKUAST-K (SC)
Salvia moocroftiana
Benth.
Lamiaceae Moorcroft’s Sage
Perennial
herb
Wild Apis, Bombus
Macroglossum
PN 3420'19.94" 7441'59.41" 1983.08 Ajas Bandipore
Sambucus wightiana
Wight and Arn.
Caprifoliaceae Kashmir Elder Perennial
herb
Wild Apis,
Lasioglossum
PL 3419'04.04" 7444'05.06" 3203.38 Gager Patri
Sanvitalia
procumbens Lam.
Asteraceae Creeping zinnia Annual herb Ornamental Apis, Bombus PL 3408'49.28" 7452'58.29" 1623.08 SKUAST-K (SC)
Saussurea costus
Lipsch.
Asteraceae Costus Kuth Perennial
herb
Wild,
Medicinal
Apis, Bombus PN 3419'47.14" 7444'05.06" 3213.33 Longmarg
Senecio
chrysanthemoides
DC.
Asteraceae Cheerful Senecio
Annual herb Wild Apis PL 341932.28" 744329.58" 2809.85 Rangnar
Sibbaldia cuneata
Kunze
Rosaceae Wedge leaf
Sibbaldia
Perennial
herb
Wild Apis PN 3419'01.74" 7445'51.88" 3252.62 Lalmarg
Sisymbrium irio L. Brassicaceae Rocket Mustard Annual herb Wild Apis PN 3408'51.04" 7452'58.97" 1622.46 SKUAST-K (SC)
Sium latijugum
Clarke
Apiaceae Squirrel tail Annual herb Wild Apis,
Sphaerophoria
PN 340859.57" 7452'57.99" 1622.46 SKUAST-K (SC)
Skimmia anquetilia Rutaceae Shrub Wild Apis, Andrena, PN 3419'28.49" 7443'36.69" 2868.92 Rangnar
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
35
Shaw Waspa,
Formica
Solanum melongena
L.
Solanaceae Bringal/ Egg
plant
Annual herb Olericulture Apis, Bombus PN 3408'50.26" 7452'44.32" 1609.23 SKUAST-K (SC)
Solanum nigrum L. Solanaceae Black Nightshade
Annual herb Wild Apis. Bombus PN 3408'53.47" 7452'58.13" 1621.23 SKUAST-K (SC)
Solanum tuberosum
L.
Solanaceae Potato Annual herb Olericulture Apis, Bombus PN 3408'52.11" 7452'47.08" 1611.69 SKUAST-K (SC)
Sonchus oleraceus
L.
Asteraceae Milk or sow
thistle
Annual herb Wild Apis,
Sphaerophoria
PL 3408'47.52" 7452'55.79" 1622.15 SKUAST-K (SC)
Sophora japonica L.
Fabaceae Tree Wild Apis, Xylocopa
PN 3407'55.50" 7450'28.65" 1602.15 Hazratbal
Stachys floccosa
Benth.
Lamiaceae Woolly
Woundwort
Perennial
herb
Wild Apis,
Polyommatus
PN 3408'49.29" 7452'54.81" 1617.54 SKUAST-K (SC)
Stellaria media Vill. Carophyllaceae Chickweed Perennial
herb
Wild Apis, Vanessa PN 3408'49.77" 7452'57.57" 1622.15 SKUAST-K (SC)
Sternbergia lutea
Spreng.
Amaryllidaceae Lily of the Field Perennial
herb
Ornamental Apis PN 3419'54.58" 7440'50.39" 1654.77 Ajas Bandipore
Syringa emodi Royle
Oleraceae Himalayan Lilac Shrub Wild Apis, Waspa
Sphaerophoria
PL 3419'59.34" 7440'56.25" 1677.23 Ajas Bandipore
Tagetes patula L. Asteraceae French Marigold Annual herb Ornamental Apis, Bombus PL 3408'48.22" 7452'56.94" 1623.08 SKUAST-K (SC)
Tanacetum vulgare
L.
Asteraceae Tancy Perennial
herb
Wild Apis, Bombus PL 3418'39.36" 744421.83" 3451.7 Gager Patri
Taraxacum
officinale Wigg.
Asteraceae Dandelion Haandh Annual herb Wild Apis,
Lasioglossum,
Scathophagia,
Halictus
PN 3408'47.01" 7452'50.26" 1613.54 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
36
Ceratina,
Andrena
Pontia, Colias.
Mylabris
Thymus linearis
Benth.
Lamiaceae Linear Leaved
thyme
Annual herb Wild Apis, Pieris,
Vanessa
NC 3410'29.71" 7454'34.17" 1826.46 Dara
Thymus serphyllum
L.
Lamiaceae Shepherd's thyme
Annual herb Wild Apis, Pieris,
Vanessa
NC 3419'53.04" 7441'08.35" 1691.38 Ajas Bandipore
Trifolium pratense
L.
Fabaceae Red Clover Annual herb Wild Apis, Bombus,
Issoria
PN 3408'48.45" 7452'51.68" 1614.46 SKUAST-K (SC)
Trifolium repens L. Fabaceae White Clover Perennial
herb
Wild Apis, Bombus,
Issoria
PN 3408'48.77" 7452'51.76" 1614.46 SKUAST-K (SC)
Tulipa stellata Hook.
Liliaceae
Himalayan White
Tulip
Annual herb Wild Apis,
Scathophagia,
Lasioglossum,
Halictus
PN 3408'50.93" 7452'56.15" 1617.84 SKUAST-K (SC)
Valeriana hardwickii
Wall.
Caprifoliaceae Indian valerian Perennial
herb
Wild Apis PN 3419'27.27" 7443'19.89" 2879.38 Rangnar
Verbascum thapsus
L.
Scrophulariaceae Great mullein Annual herb Wild Apis, Formica PN 3408'49.30" 7452'51.97" 1614.77 SKUAST-K (SC)
Veronica arvensis L.
Scrophulariaceae Speedwell Annual herb Wild Apis,
Lasioglossum,
Aulocera
PN 3408'59.42" 7452'47.81" 1613.23 SKUAST-K (SC)
Veronica persica
Poir.
Scrophulariaceae Persian
speedwell
Annual herb Wild Apis, Pieris
Polyommatus,
PN 3408'49.42" 7452'57.81" 1614.46 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
37
Aulocera
Viburnum
grandiflorum Wall.
ex DC.
Adoxaceae Grand Viburnum
Kilmesh Shrub Wild,
medicinal
Apis, Pieris,
Vanessa
PN 3440'20.78" 7443'00.64" 2426.46 Ahrabal
Viola biflora L. Violaceae Yellow wood
violet
Annual herb Wild Apis, Pieris NC 3408'45.45" 7452'36.25" 1606.77 SKUAST-K (SC)
Viola odorata L. Violaceae Wood violet Banaf sha
Annual herb Wild Pieris ,
Sphecodes
NC 3408'55.86" 7452'10.55" 1660 SKUAST-K (SC)
Weigela floribunda
C.A.Mey.
Caprifoliaceae Shrub Ornamental Apis, Neptis PN 3408'49.44" 7452'40.89" 1606.77 SKUAST-K (SC)
Zea mays L. Poaceae Maize Makai Annual herb Wild Apis PL 3408'51.41" 7452'50.43" 1614.77 SKUAST-K (SC)
Zinnia elegans L. Asteraceae Zinnia Annual herb Ornamental Apis, Bombus,
Neptis
PN 3408'48.11" 7452'58.99" 1623.69 SKUAST-K (SC)
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
38
Table 1.2: Taxonomic conspectus of insect visitors recorded during the present
study in the Kashmir Himalaya
Family Genera Species
Apidae
Xylocopa Xylocopa valga Gerstaecker, 1872
Xylocopa violacea Linnaeus 1758
Bombus Bombus simillimus Smith 1852
Bombus tunicatus Smith 1852
Bombus trifasciatus Smith1852
Bombus rufofasciatus Smith, 1852
Bombus asiaticus Morawitz, 1875
Bombus pyrosoma Morawitz 1890
Bombus miniatus Bingham, 1897
Amegilla Amegilla fallax Smith 1879
Apis Apis cerana Fabricius 1793
Apis mellifera Linnaeus 1761
Halictidae
Lasioglossum
Lasioglossum himalayense Bingham 1898
Lasioglossum nursei Bluthgen 1926
Lasioglossum rugolatum Smith 1853
Lasioglossum polyctor Bingham 1908
Lasioglossum marginatum Brulle 1832
Lasioglossum sublaterale Bluthgen 1931
Lasioglossum leucozonium Schrank 1781
Halictus Halictus constrictus Smith 1853
Halictus propinquus Smith 1853
Sphecodes
Sphecodes tantalus Nurse 1903
Sphecodes lasimensis Bluthgen 1927
Andrenidae
Andrena
Andrena patella Nurse 1903
Andrena cineraria Linnaeus 1758
Andrena floridula Smith 1878
Andrena flavipes Panzer 1799
Ceratina Ceratina hieroglyphica Smith 1854
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
39
Ceratidae Ceratina propinqua Cameron, 1897
Megachalidae
Anthidium Anthidium conciliatum Nurse, 1903
Megachile Megachile conjuncta Smith, 1853
Megachile rotundata Fabricius 1793
Heriades Heriades sp.
Tenthredinidae Athalia Athalia sp.
Vespidae Vespa Vespa velutina Lepeletier, 1836
Vespa sp.
Formacedae Formica Formica sp.
Pyrrhocoridae Pyrrhocorus Pyrrhocorus sp.
Syrphidae
Metasyrphus Metasyrphus bucculatus Rondani, 1857
Sphaerophoria
Sphaerophoria bengalensis Macquart, 1842
Sphaerophoria macrogaster Thomson, 1869
Episyrphus Episyrphus balteatus De Geer, 1776
Eristalodes Eristalodes paria Bigot 1880
Eristalis Eristalis tenax Linnaeus 1758
Eristalis cerealis Fabricius,1805
Eristalinus Eristalinus sp.
Eristalinus aeneu Scopoli, 1763
Phytomya Phytomya zonata Fabricius, 1787
Bibionidae Bibio Bibio sp.
Plecia Plecia sp.
Scathophagidae Scathophagia Scathophagia sp.
Scathophaga stercoraria Linnaeus, 1758
Muscidae Ophyra Ophyra sp.
Musca Musca domestica Linnaeus, 1758
Calliphoridae Calliphora Calliphora lata Coquillett 1898.
Hesperiidae Hesperia Hesperia comma Fabricius, 1793
Lycaenidae Polyommatus Polyommatus icarus Rottemburg, 1775
Vanessa Vanessa cashmiriensis Kollar 1848
Aulocera Aulocera padma Kollar, 1844
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
40
Nymphalidae
Issoria Issoria lathonia Linnaeus, 1758
Neptis Neptis sp.
Sphingidae Macroglossum
Macroglossum nycteris Kollar 1844
Pieridae
Pontia Pontia daplidice Linnaeus 1758
Colias Colias philodice Godart, 1819
Colias fieldi Menetries, 1855
Pieris Pieris brassicae Linnaeus 1758
Meloidae Mylabris Mylabris pustulata Thunberg 1821
Scarabaeidae Cetonia Cetonia aurata Linnaeus 1758
Coccinellidae Coccinella Coccinella magnifica Redtenbacher 1843
Thripidae Frankliniella Frankliniella sp.
Chapter –
1
Figure 1.2:
Number of species distributed in different families of the
flowering plants recorded during present study.
Figure 1.3:
Percentage contribution of different growth
plants recorded during present study.
0
5
10
15
20
25
30
35
40
No. of species
Perennial herb
32%
Shrub
12%
1
Taxonomic diversity of foraging plants and their insect visitors
41
Number of species distributed in different families of the
flowering plants recorded during present study.
Percentage contribution of different growth
-
forms in the flowering
plants recorded during present study.
Families
Biennial herb
3%
Tree
10%
Taxonomic diversity of foraging plants and their insect visitors
Number of species distributed in different families of the
forms in the flowering
Annual herb
43%
Chapter –
1
Figure 1.4:
Wild and cultivated status of the flowering plants recorded during
present
Figure 1.5:
Number and percentage of the flowering plants with reference to
insect visitor reward
nectar)
Floriculture
37, 16%
Social Forestry
4, 2%
Horticulture
15, 7%
1
Taxonomic diversity of foraging plants and their insect visitors
42
Wild and cultivated status of the flowering plants recorded during
study (number and percentage)
Number and percentage of the flowering plants with reference to
insect visitor reward
(PL=pollen, NC=nectar, PN= both pollen and
152, 67%
Social Forestry
Horticulture
15, 7%
Agriculture
19, 8%
PN
180,79%
PL
42,19%
NC
5,2%
Taxonomic diversity of foraging plants and their insect visitors
Wild and cultivated status of the flowering plants recorded during
Number and percentage of the flowering plants with reference to
(PL=pollen, NC=nectar, PN= both pollen and
Wild
152, 67%
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
43
Figure 1.6: Number of species belonging to different families of insect flower
visitors recorded during present study.
DISCUSSION
The results obtained during the present study revealed that the Kashmir
Himalaya has a rich floristic diversity interacting with diversified flower
visitors. A total of 227 plant species were found to act as feeding and nesting or
mating sites for different flower visitors, which have diverse habits (trees,
shrubs and herbs) and function as pollen and nectar source for flower visitors
(Table 1.1; Figs. 1.2-1.5) including social bees, thus the present study area is
suitable for commercial apiculture as bee flora (Paray et al., 2014; Behera et
al., 2014; Abrol et al., 1989; Singh 1982; Dukku 2013). The results
demonstrated that the Kashmir Himalaya has a rich bee flora diversity which
can be used for production of honey, wax etc. qualitatively as well as
quantitatively and can be used to boost pollination service by enhancing the
populations of pollinators (Partap 1997). The diverse pollen and nectar
0
2
4
6
8
10
12
14
No. of species
Family
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
44
producing plants found in Kashmir Himalayas as revealed by the present study
(Table 1.1) vary in habit, purpose and reward presentation as reported by other
studies as well ( see Suryanarayana et al., 1992; Ramalho et al., 1990; Momose
et al., 1998; Gaur et al., 2014). Among different foraging plant families,
Asteraceae was the largest family followed by Rosaceae and Lamiaceae (Fig.
1.2). The present study revealed that Kashmir Himalaya has a rich nectar
resource diversity which greatly helps in framing flower visitor community
structure (Potts et al., 2004).
The results from this study offer novel insights regarding further studies
on the pollination services. Furthermore, as a source of employment generation
in the region, bee keeping is the result of diversity and abundance of flowering
plants. Bee keeping largely depends on type of forage plants of a particular area
(Burkle and Irwin, 2011, Behera 2014). Pollen and nectar are the main reward
for bees during pollen transfer. These two rewards become main ingredients of
their food and for production of honey (Nnamani et al., 2013). Bee keeping
potential of an area depends upon pollen quantity and nectar volumes of plants
of that particular area (Akratanakul 1990). Bee flora quantification and
mapping shall be the future challenges for properly assessing the pollinator
fauna and sustenance of bee-keeping (Datta et al., 2008). The documented
plant species varying in rewards, prevailing pollen and nectar as a common
reward, can serve for commercial bee-keeping industry at large scale (Abrol et
al., 1989; Singh, 1982; Dukku, 2013). The key role of pollination in sustainable
ecosystemic output makes the conservation of pollinators important (Adhikari,
2003). The knowledge flow to farmer and bee-keepers about the bee flora, its
significance, role and need of conservation can increase the quality and
quantity of agricultural outcomes (Partap, 1997). Research findings revealed
that Salix species (Salix caprea and S. alba) acts as the potential source of
pollen and nectar for pollinators.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
45
The present study revealed 70 different species of flower visitors (Table
1.2) from Kashmir Himalaya that visit 227 above-documented plant species for
their feeding and nesting purposes. During the present study, majority of flower
visitor species were belonging to the order Hymenoptera, which is in
conformity with previous studies (Jasara and Rafi, 2008; Hussain et al., 2012).
Among diverse insect visitor families, Apidae was largest followed by
Halictidae (Fig. 1.6). However, Paray et al. (2014) while focussing on diversity
of insect pollinators on single plant species of apple (Malus domestica)
observed the diversity of Halicted bees surpassing other pollinators in the
Kashmir valley. Among the halictid bees, Lasioglossum genus was highly
diversified and abundant (Ganie et al., 2013; Paray et al., 2014). Abundance of
flower visitors was found at peak during the flowering period of these plant
species. Two groups of flower visitors have been recorded: bee visitors and
non-bee visitors. The honey bees, bumble bees, halicted bees, carpenter bees
constituted the bee visitors, where as the wasps, butterflies, thrips, hoverflies,
ants and moths were recognised as non-bee visitors. The present study assumes
significance by providing an inventory of plants used for nesting and feeding
by flower visitors in the Kashmir Himalaya, thus cataloguing the crucial
interaction between these two forms of life. Quite often, it has been seen that
the diversity of flower visitors is directly proportional to diversity of flowering
plants of a particular region (Nicholls and Altieri, 2012). The foraging
activities of flower visitors enhance the performance of a cross pollinated crops
in a kind of mutualistic relationship (Sahli and Conner 2007). The diverse
insect visitors reported in this study suggests that the monopoly of honey bees
as pollinators need to be revisited and the efficiency of other flower visitors can
be tested and a specific pollinator can be recommended for a particular
geographical location. Use of the recommended pollinator and its habitat
management for pollination can fulfil the need of pollinator scarcity in
horticultural crops of this Himalayan region and hopefully increase the quality
and quantity of the cross pollinated crops, such as apple.
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
46
Flower and insect visitor interaction is the most important example of
plant insect interaction, which is a key to the maintenance and sustenance of
biodiversity. The insect visitors visit the flowers for their food and in exchange
help plants by pollen transfer. Pollen transfer, commonly called pollination, is
an essential ecological service for maintenance of plant diversity.The present
study describes the diversity of the two different forms of life (flowering plants
and insect visitors) which interact in such a way that determines their
evolutionary success and serves as a base for biodiversity sustenance. The
results can guide the need of conservation of these natural interactions between
flowers and insects, which will benefit in the form of more productive natural
ecosystem. In fact, both cultural and natural ecosystems are linked directly on
this vital biotic interaction.
CONCLUSIONS
A total 227 plant species belonging to 182 genera and 58 families were
recorded, which serve as foraging plants for different flower visitors. Annual
herbaceous plants were found as abundant source of pollen and nectar for
flower visitors followed by perennial herbs, perennial shrubs, perennial trees
and finally biennial herbs. Asteraceae was found to be dominant family of
pollen and nectar producing plants in Kashmir Himalaya. Wild flora was found
to be dominant source of pollen and nectar. Majority of pollen and nectar
producing plants were found to act as the source of both pollen and nectar (bi-
dimensional) as compared to pollen or nectar only (uni-dimensional). The
study documented 70 flower visitors belonging to 43 genera and 23 families
which were found visiting 227 plants species. Family Apidae was the dominant
group of flower visitors. The results obtained during the present study can
provide a scientific platform for future research in the field of plant pollinator
interaction, pollinator food, pollen and nectar producing plants, pollination
biology, organismic interaction, diversity and co-extinction of biodiversity in
Chapter – 1 Taxonomic diversity of foraging plants and their insect visitors
47
this Himalayan region. The results can provide baseline biodiversity
information framework for conservation of plant insect interaction and can help
to cope with currently pressing problem of pollinator decline, which has huge
implications for production of different fruit crops of Kashmir, and will help in
overall maintenance and sustenance of biodiversity in Kashmir Himalaya.
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58
INTRODUCTION
Among different types of ecosystems, the flora of pollinator-dependent
ecosystems is intimately associated with floral traits they display for their
reproductive success and plant fitness. Floral traits of a particular plant species
include both attraction and reward features, which are the drivers of floral
advertisement policy for pollinator recruitment. Pollinator eligibility is
generally determined by the type and magnitude of floral traits a flowering
plant species displays (Stebbins, 1970). Floral traits affect the behaviour of
pollinators visiting a flower and helps flower to provide access to some
pollinators and exclude others (Rodriguez and Fenster, 2008). The interaction
of flowers with pollinators is the result of floral/inflorescence traits they
display and there is a huge diversity in floral traits viz. diversity in flower
colour, floral shape, display size, floral scent, flower symmetry and
inflorescence architecture. Floral traits are the main drivers of pollinator
attraction towards the floral reward including nectar, pollen and herbivory for
successful pollen transfer. The visual/emitted signal from flowers in the form
of floral traits is recognised by pollinators and guides them towards ultimate
reward. Among the vast array of floral traits, floral colour and display are
considered to be of paramount importance. The floral traits play an important
role in structuring the pollinator community of an ecosystem. Flower parts run
a coordinated program of floral advertisement by displaying a set of
coordinated floral traits, and the attracting traits coordinate in tandem with
rewarding traits for effective pollinator recruitment. In fact, the diversity in
floral traits is the key for diversification of angiosperms and their pollinators
result into diverse plant pollinator interactions (Schiestl, 2012).
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REVIEW OF LITERATURE
It was Darwin (1862) who first of all floated the idea that floral
diversification is the result of pollinator-mediated selection of floral traits and
considered it as an important evolutionary force. Stebbins (1970) stressed that
floral traits reflect the preference of pollinators among the different flowering
plant species and pollination syndrome reflects the relation of primary
pollinator with the floral traits of a particular flowering plant. Since then, a
large body of literature has reinforced that the flower traits are linked with
pollinators and a strong correlation exists between flower traits and pollinator
visitation (Conner et al., 1996; Hegland et al., 2005).
Flower pollinator interaction is the main key in structuring the global
biodiversity and its extent. Dewenter et al. (2005) found that diversity in the
quality and quantity of floral traits reflects the specialization in flower
pollinator interaction leading to rich floristic diversity all across the globe.
Faegri and van der Pijl (1979) defined floral syndrome as integrated set of
floral traits related with a particular pollinator and further remarked that the
floral traits are morphologically and ecologically linked with pollinators of
their type and floral morphology - a strong driver of pollinator attraction -
excludes some flower visitors and provides access to others which may act as
primary pollinators for a particular flowering plant species. Research studies
have revealead that 11 floral syndromes exist among animal pollinated flora
such as bat, bee, beetle, bird, butterfly, carrion fly, fly, hawk moth, moth, non-
lying mammal and wasp, which are characterised on the basis of floral traits,
such as flower architecture, reward presentation (Proctor and Yeo, 1996).
Faegri and van der pijl (1979) stated that floral advertisement reflects
behaviour of pollinator visiting a flower in order to judge the degree of
pollination syndrome. Momose et al. (1998) studied the pollination systems in
dipterocarp forest of Malaysia and found that 12 categories of pollination
systems were related with the floral traits and developed pollination syndrome
Chapter – 2 Diversity of floral traits with reference to insect visitors
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on the basis of floral traits in dipterocarp forests. According to Johnson and
Steiner (2000), broad scale comparison between floral traits and pollinators is
the effective test for floral syndrome and the floral traits show an independent
relationship with pollinator morphology especially with the size of pollinators.
Ollerton and Watts (2000) constructed floral syndromes on the basis of single
floral trait. However later on, Ollerton et al. (2009) constructed floral
syndromes on the basis of multi-floral traits and found that no plant species fall
within such pollination syndrome and principle pollinator for two third of
flowering plants cannot be predicted.
Waser (1983) found that among the floral traits advertised for pollinator
attraction, floral rewards may be considered ultimate attractants where as
flower or inflorescence architecture including colour, odour, shape and size
acts as cue for pollinators to distinguish among floral resource diversity. The
floral advertisement follows reward presentation to pollinators and the
pollinators with similar energy requirements may receive rewards from broad
spectrum of flora. In addition to visual and olfactory stimuli that a flower
advertises, tactile stimulus is one of the important factors for pollinator
behaviour at a flower (Heinrich, 1979). Dafni (1992) reported that floral
advertisement is a complex phenomenon which affects and modifies the
pollinator’s visual, olfactory and tactile abilities and floral architecture
including colour, size, shape, scent, strength and floral reward are the main
drivers to increase floral advertisement magnitude. Modification in pollinator
behaviour at a flower is the outcome of cumulative effect of stimuli that a
flower displays. Visual advertisement by flowers is the only way to
communicate with the pollinators and advertisement timing usually coincides
with the reward presentation. Lawrence et al. (2004) studied the role of
inflorescence in floral advertisement and found that floral display of
inflorescence increases the attractiveness of pollinators and thus increases
visitation rate of pollinators, and also the consistency in pollinator movement
Chapter – 2 Diversity of floral traits with reference to insect visitors
61
varies with different inflorescence architecture. Chittka and Raine (2006) found
that flowers depend on sensory signals for advertising their rewards to pollen
vectors and for this floral display colour, size and scent are the pivotal for
pollinator attraction.
Heinrich (1975) reported a significant relationship between floral reward
presentation and energetics of pollinator. Faegri and van der Pijl (1979) found
that flowering plants differ markedly in floral traits associated with both the
attraction and reward for pollinators. Willson and Bertin (1979) found that
visitation rate of pollinators are also affected by flower number and higher the
flower number more is the visitation rate and vice versa. Waser (1983) have
examined the effect of floral scent, colour and nectar on the attraction of
pollinators and found that these traits play a crucial role in floral advertisement
and increases the visitation rate of pollinators. Armbruster (1984) reported that
occurrence of diverse floral resources which reflect the diversity in pollinators
that visit them. Menzel and Backhaus (1990) found that pollinators’ visible
spectrum is the most important factor which enables the pollinator to choose a
particular colour. Waser and Chittka (1998) found that the signal complexity
exists within each of the flowers: the visual signals vary in aspects such as size,
pattern, iridescence, colour, brightness and contrast, and further the symmetry
affects the behaviour of pollinator at a flower for pollen transport. Raguso
(2004) stated that a typical flower is a multi-sensory billboard displaying a
dazzling variety of stimuli for reward presentation to pollen vector and floral
traits act as mediators of signals for pollinators. The floral traits affect the
foraging behaviour of pollinators with different floral advertisement efficacy
for attracting different types of pollinators (Fulton and Hodges, 1999; Glaettli
and Barrett, 2008). Bosch et al. (1997) studied relationship between floral traits
and their pollinators in Spain and recorded floral traits and their flower visitors
for 17 most visited plants.
Chapter – 2 Diversity of floral traits with reference to insect visitors
62
Thompson (2001) studied pollinator visitation pattern in relation with
floral design and display in generalized pollination system and found that
number of pollinator visits was positively corelated with number of open
flowers for pollinators such as bees, butterflies and hawk moths. He also found
that butterflies visited large floral displayed plants where as hawk moths were
found only such insect type having a positive co-relation between the number
of flowers visited per foraging bout and flower size. Ohashi and Yahara (2001)
found that among plant population flowers/inflorescence with larger floral
display has high visitation rate as compared to smaller displayed flowers or
inflorescences and also the bumble bee pollinators prefer flowers with larger
display as compared to smaller one. Harder et al. (2001) found that among
inflorescence architecture, inflorescence display size is one the floral trait that
attracts and increases the visitation rate of pollinators and act as strong
advertising driver for pollinator attraction as compared to solitary flowers. Bell
(1985) found that corolla size of flower affects the visitation rate of pollinators
and larger sized corolla attracts more pollinators as compared to smaller sized
one. Conner et al. (1996) studied effect of flower size and flower number on
pollinator visitation on wild radish (Raphanus raphanistrum) and found that
pollinator visitation increases by increasing the flower size and flower number
for syrphid flies, and there was increase in visitation rate of small bees by
increasing flower number and not flower size.
Lafuente et al. (2005) studied relationship between floral trait and
pollinator visitation and found that floral display is the important driver which
affects the visitation rate of pollinators. Hegland and Totland (2005) studied
relationship between species floral traits and pollinator visitation in a temperate
grassland from Norway and found that there was increase in visitation rate by
increasing the floral display, floral form and symmetry were found important
drivers of visitation rate for flies. They concluded that flower size and flower
density are important drivers of pollinator visitation rate. Guerrero et al. (2014)
Chapter – 2 Diversity of floral traits with reference to insect visitors
63
concluded that floral traits of tropical flowering plants are specialized for
certain effective pollinators known as principal or primary pollinators as
compared to floral traits meant for general pollinators, including primary as
well as secondary. Lazaro et al. (2009) found that the floral neighbourhood
affects the level to which floral traits are related with pollinators and thus
affects the generalization-specialization continuum of flower pollinator
interaction. Petanidou and Lambron (2005) reported that a vast majority of
flower traits basically advertise for all pollinators constituting the generalized
flower pollinator interaction. Fenster et al. (2006) studied the role of floral
traits in attracting primary and secondary pollinators and found that nectar
volume, nectar sugar concentration, and total sugar reward were positively
correlated to flower size. Buide (2005) studied floral trait variation and
pollinator attraction in Silene acutifolia and found that pollinator visitation
frequency was more in plants having more open flowers as compared to less
open flowers, thus increased the seed set of more open flowered plant. Vieyra
et al. (2006) studied floral trait relation with pollinator visitation rate in
Cyclopogon elatus and found that floral traits affect pollination in multiple
ways, for instance increase in flower number increases visitation rate and
nectar depth increases pollination efficacy.
Navarro et al. (2007) studied adaptation of floral traits and mating
systems to pollinator unpredictability in Disterigma stereophyllum, a memeder
of Ericaceae from Colombia and found that floral design including a set of
different floral traits and floral rewards was intermediate to bee-pollinated and
bird-pollinated plants. Gomez et al. (2008) found that there is the strong
corelation between floral traits and floral rewards and found that pollinator may
act as a selective agent on floral traits. Olesen and Jordano (2002) found that
flower pollinator interaction in community can be visualised as a network,
where one type of floral resource with one set of floral traits is generally visited
by a variety of pollinators and each type of pollinator visits a variety of floral
Chapter – 2 Diversity of floral traits with reference to insect visitors
64
resource. Thomson (2003) found that even though flowers of a particular plant
species are qualified by different types of flower visitors with their floral traits
but their pollination efficacy varies which should be the basis for categorizing
them into primary or principle pollinators and secondary pollinators of a
particular plant species. Irwin et al. (2004) found that pollinator foraging plants
develop nectar robber-excluding traits that aid in effective pollination. Karron
et al. (2004) found that floral traits are responsible to reproductive success of
plant by increasing the visitation frequency of pollinators for a particular plant.
Grindeland et al. (2005) studied the effect of flower display and density
on pollinator visitation rate in Digitalis purpurea in Norway and found that
pollinator visitation rate gets modified qualitatively as well as quantitatively by
flower display size and density. They found increase in pollinator visitation
with increase in flower display size and dense-flowered patches. Ashman et al.,
(2005) found that pollinators approach to larger floral rewards (pollen and
nectar) as compared to lesser floral rewards. They also found that preference of
pollinators towards larger petals and corolla tubes with wider diameters. Ishii et
al. (2008) found that bumble bee forages large displayed flowers. Campbell et
al. (2010) studied the relation between flower colour and pollinator visitation
rate and found that pollinators visit yellow-coloured flowers as compared to
white-coloured flowers. Malerba and Nattero (2011) found that pollinators get
attracted towards the bunch of flowers as compared to individual ones.
Chittka and Schurkens (2001) found that flower pollinator communities
are considered as biological markets, where pollinators choose reward
according to the floral advertisement they prevail. Fabbro and Korner (2004)
studied altitudinal differences in flower traits in Switzerland and found that
flower display of both sites was similar but flowering period of higher altitude
remains for longer durations. Herrera and Pellmyr (2002) stated that plant
insect interactions may be positive or negative: herbivory and florivory are the
negative interactions, where as pollination and seed dispersal is the positive
Chapter – 2 Diversity of floral traits with reference to insect visitors
65
interactions. Schlumpberger et al. (2009) studied variation in floral characters
and its consequences for pollinator attraction in Northern Argentina and found
that sphingid pollination is prevalent only in populations with the longest
flowers, in which floral morphological traits and nectar volumes match the
expectations for the sphingid moth, thus constituting hawk moth pollination
syndrome. Bastolla et al. (2009) found that specialization in flower pollinator
interaction reduces the competition among pollinators and increases their
diversity index.
Andrews et al. (2007) studied the attraction of pollinators and
herbivores to floral volatiles in Cucurbita and found that the plant emits three
different types of volatile chemicals: one attracts only pollinators, second one
attracts only herbivores, where as third attracts both. Howell and Alarcon
(2007) found that pollinators, especially bees are able to recognise reward by
the scent of a particular plant species, bees learn odour of a flower faster and
remember it for longer and pollen odour play vital role in recognition of plant.
Raguso (2008) found that flower scent vary in composition, abundance,
intensity and ratio that modify the pollinator behaviour and promotes
specialization in flower pollinator interactions. Majetic et al. (2009) found that
floral scent acts to increase pollinator attraction for a particular flowering plant
species, thus improving the reproductive success by optimising pollen import
and export by pollen vector. Waelti et al. (2009) studied pollinator
attractiveness in dioecious plants of Silene latifolia and found that male plants
produced more flower with higher content of floral scent as compared to
female flowers with the pollinator-moth, Hadena bicruris. Kessler et al. (2012)
studied floral trait relationship with pollinator attraction and found that there
are some floral traits like floral scent which helps flower in fair recruitment of
pollinator and avoiding attack from florivores, because both mutualists as well
as antagonists are eligible to floral recruitment policy. Morales et al. (2013)
studied the role of sterile flowers in pollinator attraction and reproductive
Chapter – 2 Diversity of floral traits with reference to insect visitors
66
success and found that presence of sterile flowers within fertile flowers
increases the quality and quantity of pollination by promoting pollen import
and export. Filella et al. (2013) studied role of floral scent in changing flower
pollinator market and found that scent advertisement is higher when there is
competition for pollinator attraction in plentiful floral resource. They also
found that less abundant flowering species compete with dominant species by
advertising scent. Vega et al. (2014) studied role of floral scent in ant
pollination and found that floral scent acts as a floral trait that plays an
important role in ant attraction to a flower.
MATERIALS AND METHODS
During the present study, observations in the field and laboratory were
conducted in different months of the year 2014-2015 in the study area to record
and measure the floral traits. The floral traits are characters displayed by
flowers in order to attract insect visitors. Nine different floral traits were
investigated during the present study which include inflorescence architecture
(or inflorescence type), flower/inflorescence display size, flower reward,
flower shape, flower symmetry, flower sex, flower chromatism, corolla colour,
anther colour and pollen colour with the help of naked eye for 215 foraging
plant species (Willson and Bertin, 1979, Conner et al., 1996; Grindeland et al.,
2005; Hegland and Totland 2005). Flower scent was observed by smelling the
host plant species.
(i) Inflorescence architecture
Inflorescence architecture includes infloresce type which refers to
arrangement of flowers on the floral axis. Based on the type of inflorescence,
the foraging plants were grouped into 06 categories: raceme, cymose,
capitulum, catkin, glomerule and cyathium.
Chapter – 2 Diversity of floral traits with reference to insect visitors
67
(ii) Flower/Inflorescence display
Flower/ inflorescence display of foraging plants is the plateform for
flower visitors and its size was investigated by measuring the edge to edge
distance of a flower/inflorescence with the help of a measuring scale (Buide,
2004). The measurement of floral display size was carried out for plants
displaying flowers as advertising unit, and the inflorescence display size was
measured for plants either having very small flower display or displaying
inflorescences as a unit of advertisement. During the present study, the floral
display size measurement of plants displaying flowers as a unit of
advertisement, revealed a great variability in the floral display size ranging
from 0.5 cm - 12 cm, and has been classified into 5 categories (Table 2.1):
Very small floral display (VSFD), Small floral display (SFD), Medium floral
display (MFD), Large floral display (LFD) and Very large floral display
(VLFD) on the basis of size of floral display. If flower size found to be very
small, then inflorescence display size was taken as reference. Inflorescence
display size was also categorized into 5 categories as Very small inflorescence
display (VSID), Small inflorescence display (SID), Medium inflorescence
display (MID), Large inflorescence display (LID) and Very large inflorescence
display (VLID) on the basis of size of inflorescence display (Table 2.2).
Table 2.1: Categories of floral display size
Category of floral display Lower range
(cm)
Upper range
(cm)
Very small floral display (VSFD) 0.1 1.00
Small floral display (SFD) 1.1 3.00
Medium floral display (MFD) 3.1 8.00
Large floral display (LFD) 8.1 10.00
Very large floral display (VLFD) 10.1 12.00
Chapter – 2 Diversity of floral traits with reference to insect visitors
68
Table 2.2: Categories of inflorescence display size
Category of inflorescence display Initial
range (cm)
Final Range
(cm)
Very small inflorescence display (VSID) 1.00 5.00
Small inflorescence display (SID) 5.00 10.00
Medium inflorescence display (MID) 10.00 20.00
Large inflorescence display (LID) 20.00 25.00
Very large inflorescence display (VLID) 25.00 30.00
(iii) Flower reward
Flower rewards the flower visitors for visiting them in the form of
pollen and nectar. Flower reward an insect visitor receives was observed by the
foraging behaviour of insect visitor at the flower. The dancing foraging
behaviour of an insect visitor over the flowers/anthers revealed the pollen
presentation of host flower; where as sucking foraging behaviour of an insect
visitor usually at the base of petal revealed nectar presentation by the flower;
and an insect visitor showing both these behaviours revealed the dual
presentation from the flower as a reward. Thus the foraging plants were
classified into three groups on the basis of reward presentation: pollen and
nectar rewarding plants (PN), pollen rewarding plants (PL) and nectar
rewarding plants (NC).
(iv) Flower shape
Flower shape includes the shape of perianth, calyx, corolla or
hypanthium (if present). The various forms of flower shape shown by the
foraging plants were classified into 17 categories: rotate, ligulate and tubular,
bilabiate, saucer, bowl, campanulate, cruciform tubular, trumpet, rosaceous,
Chapter – 2 Diversity of floral traits with reference to insect visitors
69
infundibular, personate, paplionaceous, cup, ligulate, coronate and
caryphylaceous. Each of these flower shapes defines the particular set of floral
trait.
(v) Flower symmetry
The flower symmetry was observed by keen observation of flowers in
the field. Two categories were recognised: flowers with radial symmetry were
classified as actinomorphic and flowers with bilateral symmetry or
asymmetrical shape were classified as zygomorphic flowers (Raguso, 2004;
Guerrero et al., 2014).
(vi) Flower sex
Plants bearing flowers with only one type of sex organs were grouped as
unisexual flowers that were further classified into male unisexual flowers
(stigmatic) and female unisexual flowers (pistillate), and plants bearing flowers
with two different sexes in the same flower were classified as bisexual flowers.
(vii) Flower chromatism
The flower chromatism is the cumulative colour a flower/inflorescence
displays. The flowers/inflorescences which showcased only one colour from
their display were grouped as monochromatic flowers; flowers which displayed
two colours either co-dominantly or in mixed form were classified as
dichromatic flowers; and the flowers which displayed multiple colours from the
same floral display or different floral displays of same species were classified
as multichromatic flowers.
(viii) Flower colour
The flower colour is the colour of the various parts of a flower such as
perianth, calyx, corolla, anther and pollen. In the present study perianth and
calyx were taken with corolla colour. On the basis of colour of the corolla tube,
the foraging plants were grouped into 27 categories: yellow, pink , blue, purple,
Chapter – 2 Diversity of floral traits with reference to insect visitors
70
cream, red, pink and white, white and yellow, violet, green, white and purple,
yellow green, red yellow and white, blue, yellow and white, pink, yellow and
white, white and violet, yellow, violet and white, red and black, blue, white and
purple, yellow and black, yellow and red, yellow green and pink, maroon, red
and white, orange, yellow and white (Campbell et al., 2010). The foraging
plants were also grouped into 20 categories on the basis of their anther colour.
Pertinently, anther colour within the floral display provides clues about the
pollinator reward. Similarly, the foraging plants were grouped into 17
categories on the basis of pollen colour they possess (Campbell et al., 2010).
RESULTS
During the present study, nine different types of floral traits were
observed for 215 foraging plant species (Table 2.3 and photoplates 2.1-2.3).
Chapter – 2 Diversity of floral traits with reference to insect visitors
71
Out of them the first one inflorescence architecture was investigated in
215 plant species. The results revealed that majority of the plant species had
raceme type of inflorescence (98 spp.), followed by cymose (77 spp.),
capitulum (32 spp.), catkin (4 spp.), Glomerule and cyathium (2 spp. each)
(Fig. 2.1).
Figure 2.1: Contribution of different inflorescence types in the pollinator
flora.
Among the total of 215 plant species, flower display size was measured
for 155 plants where as inflorescence display size was measured for 60 plant
species. The results revealed that largest number of plant species (71 spp.)
belonged to SFD category, followed by MFD (42 spp.), VSFD (26 spp.),
VLFD (07 spp.) and LFD (06 spp.) (Table 2.3, Fig. 2.2)
0
20
40
60
80
100
120
Types of inflorescence
No. of species
Chapter – 2 Diversity of floral traits with reference to insect visitors
72
Figure 2.2: Number of species in different floral display size classes.
The results revealed that among different inflorescence display size
classes, VSIDwas the largest (27 spp.), followed by SID (19 spp.), MID (11
spp.), LID (02 spp.) and finally VLID display (01 sp.) (Table 2.1, Fig. 2.
Figure 2.3: Number of species belonging to different inflorescence display
size classes.
0
10
20
30
40
50
60
70
80
VSFD SFD MFD LFD VLFD
No. of species
Floral display size classes
0
5
10
15
20
25
30
VSID SID MID LID VLID
Inflorescence display size classes
No. of species
Chapter – 2 Diversity of floral traits with reference to insect visitors
73
The results for the floral reward presentation revealed that, among the
215 plant species investigated, 161species present reward of both pollen and
nectar, 44 species pollen only, and the remaining 10 species nectar only (Fig.
2.4 and photoplates 2.1-2.3).
Figure 2.4: Distribution of flower reward type in terms of number of species
and percentage among pollinator flora.
Majority of insect-visited flora have rotate form of flowers (38 spp.)
followed by bilabiate (29 spp.), ligulate and tubular (25 spp.), saucer (22 spp.),
bowl (21 spp.), campanulate (15 spp.), cruciform (11 spp.), trumpet (10 spp.),
tubular (09 spp.), rosaceous (7 spp.), infundibular, personate, paplionaceous
(05 spp. each), cup, ligulate (04 spp. each), coronate (03 spp.) and
caryphyllaceous (01) (Fig. 2.5 and photoplates 2.1-2.3).
PN
161,75%
PL
44,20%
NC
10,5%
Chapter – 2 Diversity of floral traits with reference to insect visitors
74
Figure 2.5: Number of species belonging to different floral shapes recorded in
the pollinator flora of Kashmir Himalaya.
The results on the floral trait of flower symmetry revealed that, among
the 215 plant species studied, 152 plants species possessed actinomorphic
flowers, 44 species had zygomorphic flowers and 19 species showed both the
types of flower symmetry (Fig. 2.6 and photoplates 2.1-2.3).
Figure 2.6: Distribution of flower symmetry in terms of number of species and
percentage among the pollinator flora.
0
5
10
15
20
25
30
35
40
No. of species
Flower shape
Actinomorphic
152,71%
Zygomorphic
44,20%
Both
19,9%
Chapter – 2 Diversity of floral traits with reference to insect visitors
75
On investigation of flower sex among 215 plant species, it was revealed
that 208 species were bisexual, where as 07 species were unisexual (Fig. 2.7).
Figure 2.7: Distribution of floral sex in terms of species richness and
percentage among the pollinator flora.
The results of the flower chromatism revealed that among 215 plants
investigated, 169 plant species showed monochromatism, followed by
dichromatism (35 spp.) and multichromatism (11 spp.) (Fig. 2.8).
Figure 2.8: Distribution of flower chromatism in terms of number of species
and percentage among the pollinator flora.
Bisexual
208, 97%
Unisexual
7, 3%
Monochromatism
169,79%
Dichromatism
35,16%
Multichromatism
11,5%
Chapter – 2 Diversity of floral traits with reference to insect visitors
76
The results revealed diverse forms of corolla colours in the 215 plant
species investigated. The yellow colour flowers dominanted with 52 species,
closely followed by white (49 spp.), and then pink (25 spp.), blue (21 spp.),
purple (18 spp.), cream, red (06 spp. each), pink and white, white and yellow,
(05 spp. each), violet, green (04 spp. each), white and purple (03 spp.), yellow
green (02 spp.), red yellow and white (02 spp. each) and remaining blue,
yellow and white, pink, yellow and white, white and violet, yellow, violet and
white, red and black, blue, white and purple, yellow and black, yellow and red,
yellow green and pink, maroon, red and white, orange, yellow and white
represented by one species each (Fig. 2.9 and photoplates 2.1-2.3).
Figure 2.9: Number of species belonging to different corolla colour types in
the pollinator flora of Kashmir Himalaya.
Similarly, diverse forms of anther colours were recorded among the 214
plant species investigated. The yellow anther colour was predominant
possessed by 120 plant species, followed by white (44 spp.), cream (12 spp.),
golden (6 spp.), purple (5 spp.), violet (5 spp.), pink (4 spp.), orange (3 spp.),
0
10
20
30
40
50
60
Corolla colour
No. of species
Chapter – 2 Diversity of floral traits with reference to insect visitors
77
dark golden (3 spp.), blue (2 spp.), yellow green (2 spp.), and the remaining
anther colours of maroon, green, brick, gray, black, mandhi (dark green), red,
golden brown and brown with one species each (Fig. 2.10 and photoplates
2.1-2.3).
Figure 2.10: Number of species belonging to different anther colour types in
the pollinator flora of Kashmir Himalaya.
On examination of pollen colour, it was revealed that among the 215
plant species, 121 species had yellow coloured pollen followed by white (50
spp.), cream (18 spp.), pink, golden (04 spp. each), dark golden and orange (03
spp. each), purple, black (02 spp. each) and others such as mandhi, red, black,
brick, blue, gray, brown, green, golden brown by one species each (Fig. 2.11
and photoplates 2.1-2.3).
0
20
40
60
80
100
120
140
No. of speciezs
Anther colour
Chapter – 2 Diversity of floral traits with reference to insect visitors
78
Figure 2.11: Number of species belonging to different pollen colour types in
the pollinator flora of Kashmir Himalaya.
Among the 215 plant species investigated, the floral scent was found to
intensify advertisement efficiency in 31 species, while as in rest of the plant
species scent presentation was lacking (Fig. 2.12).
Figure 2.12: Presence and absence of flower scent among the pollinator flora.
0
20
40
60
80
100
120
140
No. of species
Pollen colour
Absent
184,86%
Present
31,14%
Chapter – 2 Diversity of floral traits with reference to insect visitors
79
Table 2.3: Floral trait database recorded in the pollinator flora of Kashmir Himalaya; (*Abbreviation used: PN = Pollen and nectar producing
plants, PL = Pollen producing plants, NC = Nectar producing plants, AC= Actinomorphic, ZY= Zygomorphic, HM= Hermaphrodite,
UN= Unisexual, MC= Monochromatic, DC= Dichromatic, MAC= Multichromatic, A= Absent, P= Present, F= Flower,
IF=Inflorescence).
Species Infloresence
type Flower shape Flower
symmetry
Plant
sex
Flower
chrom-
atism
Corolla
colour
Anther
colour
Pollen
colour
Flower
scent
Flower
reward
Flower display
size/
Inflorescence
length (cm)
Pollinator
visits/10
min.
Abelia grandiflora
Rehd. Cymose Campanulate AC HM MC White White White A PN F 1.5 2
Abelmoschus
esculentus Moench Solitary Bowl AC HM DC Cream Maroon Cream A PN F 9 5
Aconitum laeve
Royle Raceme Hooded ZY HM DC White and
Purple Purple Purple A PN F 3.5 2
Aconogonum molle
Hara Panicle Rotate AC HM MC White White White A PL IF 14 4
Actinidia deliciosa
Liang and Ferguson.
Cymose Cup AC UN MC White Golden Cream A PN F 5 4
Aesculus indica
Hook. Panicle Saccate ZY HM DC White Pink Pink A PN F 3.2 3
Ageratum
houstonianum Mill. Capitulum tubular AC HM MC Purple Purple Cream A PL IF 3 4
Agrimonia pilosa
Ledeb. Spike rotate AC HM MC Yellow Yellow Yellow A PN F 0.8 2
Ailanthus altissima Panicle rotate AC UN MC Green Golden Cream P PL F 1.2 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
80
Swingle
Albizia julibrissin
Durazz. Raceme tubular AC HM DC
White and
Pink Yellow Yellow A PL F 4.5 3
Alcea rosea L. Spike Saucer AC HM MC Pink White White A PN F 10 6
Allium cepa L. Umbel rotate AC HM MC White Yellow Yellow P PL IF 6 6
Allium sativum L. Umbel rotate AC HM MC White Yellow Yellow P PL IF 6 6
Anaphalis busua
DC. Corymb Tubular AC HM DC
White and
Cream Yellow Cream P PL F 10 3
Anemone obtusiloba
Don Cymose rotate AC HM DC Blue Yellow Yellow A PN F 4.5 1
Anemone
tetrasepala Royle Cymose rotate AC HM DC
White and
Yellow Yellow Yellow A PN F 4.5 1
Anthemis cotula L. Capitulum Ligulate and
Tubular Both HM DC
White and
Yellow Yellow Yellow P PL IF 3 3
Antirrhinum majus
L. Raceme Personate ZY HM MAC
Red,
Yellow,
White
Yellow Yellow A PN F 4.5 3
Aquilegia fragrans
Benth. Solitary Spurred AC HM MAC White,
Purple Golden Golden A PN F 4 4
Arabis glabra
Bernh. Corymb cruciform AC HM MC White Yellow Yellow A PN F 0.6 2
Arctium lappa L. Capitulum tubular AC HM MC Purple Purple Purple A PL IF 2.5 2
Aster thomsonii
Clarke Capitulum Ligulate and
Tubular Both HM MAC
Blue,
white and
Yellow
Yellow Yellow A PN IF 3 2
Astragalus Raceme Paplionaceous
ZY HM MC Yellow Yellow Yellow A PN F 3 4
Chapter – 2 Diversity of floral traits with reference to insect visitors
81
grahamianus Benth.
Bellis perennis L. Capitulum Ligulate and
Tubular Both HM DC
White and
Yellow Yellow Yellow P PN IF 2.5 3
Berberis lycium
Royle Raceme Cup AC HM MC Yellow Yellow Yellow P PN F 0.6 2
Bergenia ligulata
Engl. Corymb Campanulate AC HM MC Cream Yellow Yellow A PN F 2 2
Bidens tripartita L.
Capitulum tubular AC HM MC Yellow Yellow Yellow A PL F 0.5 1
Brassica compestris
Corymb cruciform AC HM MC Yellow Yellow Yellow A PN F 1.4 3
Brassica rapa L. Corymb cruciform AC HM MC Yellow Yellow Yellow A PN F 1.6 3
Buddleja davidii
Franch. Panicle Salverform AC HM DC Purple Orange Orange A PL IF 20 7
Calendula
officinalis L. Capitulum Ligulate and
Tubular Both HM DC Purple Yellow Yellow A PN IF 11 5
Caltha alba
Cambess. Corymb rotate AC HM DC
White and
Yellow Yellow Yellow A PN F 4 3
Campanula
cashmeriana Royle Spike Campanulate AC HM MC Purple Cream Cream A PN F 2.4 2
Campanula
rotundifolia L. Spike Campanulate AC HM MC Blue Cream Cream A PN F 2.6 2
Campsis grandiflora
Schum. Panicle trumpet AC HM MC Red Golden Golden A NC F 4 4
Capsella bursa-
pastoris Corymb cruciform AC HM MC White Cream Cream A PN IF 2.5 1
Carduus edelbergii
Rech.f. Capitulum Ligulate and
tubular AC HM MC Purple White White A PN IF 3.5 3
Chapter – 2 Diversity of floral traits with reference to insect visitors
82
Carya illinoinensis
Koch Catkin Catkin AC UN MC Green Green Green A PL IF 12 4
Castanea sativa
Mill. Catkin Catkin AC UN MC Cream Cream Cream A PL IF 16 8
Centaurea cyanus
L. Capitulum Ligulate and
tubular AC HM MC Blue White White A PL IF 6 3
Centaurea iberica
Spreng. Capitulum Ligulate and
tubular AC HM DC
White and
Violet White White A PN IF 3.5 2
Cercis canadensis
L. Raceme Bilabiate ZY HM DC Pink Golden
Brown
Golden
Brown A NC F 3 3
Chenopodium
album L. Panicle Globose AC HM MC Green Dark
golden
Dark
golden A PL IF 11 2
Chrysanthemum
coronarium Spach Corymb Ligulate and
Tubular Both HM MAC
White,
Yellow,
Pink
Orange Orange A PN IF 7 5
Cichorium intybus
L. Capitulum ligulate AC HM MC Purple White White A PN IF 3.5 2
Cirsium arvense
Scop. Capitulum Ligulate and
tubular AC HM MC Purple White White A PN IF 3 3
Cirsium falconeri
Petr. Capitulum Ligulate and
tubular AC HM MC White White White A PN IF 4 3
Cirsium vulgare
Ten. Capitulum Ligulate and
tubular AC HM MC Violet White White A PN IF 4 3
Clematis montana
DC. Cymose rotate AC HM DC
White and
Yellow Yellow Yellow P PN F 4.5 1
Codonopsis ovata Solitary Campanulate AC HM MC Blue Yellow Yellow P PN F 2.5 1
Chapter – 2 Diversity of floral traits with reference to insect visitors
83
Benth.
Colchicum luteum
L. Solitary rotate AC HM MC Yellow Yellow Yellow A PN F 5 4
Conium maculatum
L. Umbel rotate AC HM MC White White White P PL IF 13 5
Convolvulus
arvense L. Cymose Infundibular AC HM DC
White and
pink White White A PN F 4 2
Coriandrum sativum
L. Umbel Irregular ZY HM MC White White White P PL IF 7 5
Crepis tectorum L. Capitulum Ligulate and
tubular Both HM MC Yellow Yellow Yellow A PL IF 2.5 2
Cucumis melo L. Solitary rotate AC HM MC Yellow Yellow Yellow A PN F 10 2
Cucumis sativus L. Solitary Rotate AC HM MC Yellow Yellow Yellow A PN F 3.5 2
Cucurbita maxima
Duchesne Solitary Campanulate AC HM MC Yellow Yellow Yellow A PN F 12 13
Cucurbita pepo L. Solitary Campanulate AC HM MC Yellow Yellow Yellow A PN F 10 5
Cydonia oblonga
Miller Raceme Bowl AC HM MC Pink Cream Cream A PN F 4 4
Cynodon dactylon
Pers. Spike Spike ZY HM MC Purple Golden Golden A PL IF 6 1
Cynoglossum
glochidiatus Wall Cymose Bowl AC HM MC Blue Yellow Yellow A PN F 0.5 2
Datura stramonium
L. Cymose trumpet AC HM DC White White White A PN F 2.3 2
Daucus carota L. Umbel rotate AC HM MC White Violet White A PL IF 10 5
Delphinium roylei Raceme Spurred ZY HM MC blue Yellow Yellow A PN F 2 1
Chapter – 2 Diversity of floral traits with reference to insect visitors
84
Munz
Descurainia sophia
Prantl Raceme Cruciform AC HM MC Yellow Yellow Yellow A PN F 0.6 1
Digitalis purpurea
L. Raceme Campanulate ZY HM MC Yellow Yellow Yellow A PN F 1.7 2
Dipsacus inermis
Wall. Capitulum Trumpet AC HM MC White Cream Cream A PN IF 5 4
Doronicum
falconeri
Hook. Panicle Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PN F 2.5 2
Echium
plantagineum L. Cymose Campanulate AC HM MC Blue Blue Blue A PN F 2 2
Epilobium hirsutum
L. Raceme Saucer AC HM MC Pink White White A PN F 2.2 2
Epilobium
parviflorum Schreb.
Raceme Saucer AC HM MC Purple White White A PN F 2.3 2
Epilobium
royleanum Hausskn.
Raceme Saucer AC HM MC Pink White White A PN F 2.5 2
Eremurus
himalaicus Baker. Raceme rotate AC HM MC White Brick Brick A PN F 2.5 2
Eriobotyria
japonica
Lindl. Raceme Pear AC HM MC White Yellow Yellow P PN F 2.4 3
Erysimum
hieraciifolium L. Corymb cruciform AC HM MC Orange Yellow Yellow P PN F 1 1
Eschscholzia
californica Cham. Solitary Saucer AC HM MAC
Orange,
Yellow,
White
Yellow Yellow A PN F 5 5
Chapter – 2 Diversity of floral traits with reference to insect visitors
85
Euonymus
hamiltonianus Wall.
Cymose cruciform AC HM MC Red Yellow yellow A PN F 0.5 1
Euphorbia
helioscopia L. Cyathium Bowl AC HM MC Yellow
green
Yellow
green Yellow P PN F 1.2 1
Euphorbia wallichii
Hook.f. Cyathium Bowl AC HM MC Yellow
green
Yellow
green yellow P PN F 0.7 1
Foeniculum vulgare
Mill. Umbel Disc AC HM MC Yellow Yellow Yellow P PN IF 7 8
Forsythia
viridissima Lindl. Panicle Bilabiate ZY HM MC Yellow Yellow Yellow P PN F 2.5 2
Fragaria ananassa
Duchesne Cymose Bowl AC HM MC White Pink Pink A PN F 2.3 3
Fragaria nubicola
Lacaita Cymose Bowl AC HM DC White Yellow Yellow A PN F 2.2 3
Fumaria indica
Pugsley Raceme tubular AC HM MC Pink Yellow Yellow A NC F 0.2 1
Galinsoga
parviflora Cav. Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PL F 0.3 1
Gentiana
cachemirica Decne.
Cymose Campanulate AC HM MC Blue White White A PN F 1.2 2
Geranium nepalense
Sweet Cymose Saucer AC HM MC White Violet White A PN F 1.5 1
Geranium pratense
L Cymose Saucer AC HM MC Purple Purple White A PN F 3.2 2
Geranium
wallichianum Don Cymose Saucer AC HM MC Purple Purple White A PN F 3 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
86
Geum roylei Wall. Cymose Bowl AC HM MC Yellow Yellow Yellow A PN F 1.7 1
Gladiolus
hortulanus Bailey Spike Campanulate ZY HM MC Pink White White A PN F 7 3
Hackelia uncinata
Fisch. Cymose rotate AC HM MC Blue Yellow Yellow A PN IF 2.5 2
Halianthus annus L.
Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PN IF 11 7
Hibiscus rosa-
sinensis L. Cymose Saucer AC HM MC Pink White White A PN F 12 6
Hypericum
hookerianum Wight
and Arn.
Cymose Bowl AC HM MC Yellow Yellow Yellow A PN F 4.5 5
Hypericum
perforatum L. Cymose Rotate AC HM MC Yellow Cream Cream A PL F 3.5 2
Iberis amara L. Corymb cruciform AC HM DC White Yellow Yellow A PN F 1.3 3
Impatiens
glandulifera Royle Raceme Bilabiate ZY HM MAC
Yellow,
Violet,
White
Yellow Yellow A PN F 2.1 2
Indigofera
heterantha Wall Raceme Bilabiate ZY HM MC Pink Yellow Yellow A PN F 1.1 1
Inula royleana
Clark Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PN IF 9 7
Inula racemosa
Hook. Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PN IF 10.5 7
Ipomoea tricolor
Cav. Cymose tubular AC HM MC Blue White White A PN F 3.5 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
87
Iris decora Wall. Solitary rotate ZY HM DC
White and
Purple Yellow Yellow A PN F 7 4
Iris hookeriana
Foster Solitary rotate ZY HM DC
White and
Purple Yellow Yellow A PN F 9 4
Lactuca
dolichophylla
Kitam.
Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PL IF 2 1
Lagotis
cashmeriana Rupr. Spike Bilabiate ZY HM MC Blue Blue White A PN F 1.2 1
Lamium album L. Verticilaster Bilabiate ZY HM MC White Yellow Yellow A PN F 1.5 1
Lavandula
officinalis L. Verticilaster Trumpet AC HM MC Blue yellow yellow A PL IF 11 6
Lavatera
cashmiriana
Cambess.
Solitary Saucer AC HM MC Red White White A PN F 10.5 6
Leonurus cardiaca
L. Verticilaster Bilabiate ZY HM MC White Yellow Yellow A PN F 0.6 1
Ligularia fischeri
Turcz. Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PN F 1.8 2
Lindelofia
longiflora Baill. Spike Hat AC HM MC Blue Yellow Yellow A PN F 1.4 2
Lonicera japonica
Thunb. Spike Bilabiate ZY HM MAC Yellow Yellow Yellow P NC F 2.8 4
Lotus corniculatus
L. Umbel Bilabiate AC HM MC Yellow Yellow Yellow A PN F 1.3 1
Lycopersicon Raceme rotate AC HM MC Yellow yellow Yellow A PN F 2.9 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
88
esculentum Mill.
Magnolia
grandiflora L. Solitary Bowl AC HM MC White White White P PN F 9.5 5
Mahonia borealis
Takeda Raceme Bowl AC HM MC Yellow Yellow Yellow P PN F 0.6 3
Malus domestica
Borkh. Cymose Bowl AC HM MC White Cream Cream A PN F 2.5 3
Malva neglecta
Wallr. Fascicle Campanulate AC HM MC Pink White White A PN F 2.2 2
Malva sylvestris L. Fascicle Saucer AC HM MC Pink White White A PN F 4 3
Marrubium vulgare
L Verticilaster Bilabiate ZY HM MC White Yellow Yellow A PN F 1.3 1
Meconopsis latifolia
Prain Raceme Bowl AC HM MC Blue Golden Golden A PN F 3.5 3
Medicago lupulina
L. Raceme Bilabiate ZY HM MC Yellow Yellow Yellow A PL F 0.8 1
Mentha arvensis L. Verticilaster Funnelform AC HM MC Pink Light
Gray
Light
Gray P PL IF 5 5
Mentha longifolia L.
Verticilaster Funnelform AC HM MC Violet Violet White P PL IF 6 5
Morino longifolia
DC. Verticilaster Bilabiate ZY HM MC White Yellow Yellow A NC F 0.8 1
Myosotis arvensis L.
Cincinnus Salverform AC HM MC Blue Yellow Yellow A PN F 0.6 1
Myosotis caespitosa
Schultz Cincinnus Salverform AC HM MC Blue Yellow Yellow A PN F 0.6 1
Nepeta erecta
Benth. Verticilaster Bilabiate ZY HM MC Blue Yellow Yellow P PN F 1.6 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
89
Narcissus poeticus
L. Scapose Coronate AC HM DC
White and
Yellow Yellow Yellow A PN F 3.6 4
Nasturtium
officinale Aiton Raceme cruciform AC HM MC White Yellow yellow A PN F 0.7 2
Nelumbo nucifera
Gaertn. Solitary Bowl AC HM MC Pink Yellow Yellow A PN F 11 5
Nymphaea alba L. Solitary Bowl AC HM DC White Yellow Yellow A PN F 8 4
Nymphoides peltata
Kuntze Solitary Bowl AC HM MC Yellow Yellow Yellow A PN F 2.8 2
Oxalis corniculatus
L. Umbel Trumpet AC HM MC Yellow Yellow Yellow A PN F 1.1 2
Paparver dubium L.
Solitary Cup AC HM DC Red
Black Black Black A PN F 8.5 4
Papaver
somniferum L. Solitary Cup AC HM MAC
Red,
Yellow,
White
Mandi Mandi A PN F 9.3 5
Parthenium
hysterophorus L. Capitulum tubular AC HM MC White White White A PL IF 0.5 1
Petunia alba
Ferguson andOttley Solitary Infundibular AC HM MAC
Blue,
white and
purple
White White A PN F 3.4 2
Phaseolus vulgaris
L. Raceme Bilabiate ZY HM MC Red Yellow Yellow A PN F 2.7 3
Phlomis bracteosa
Benth. Verticilaster Bilabiate ZY HM MC Pink Yellow Yellow A PN F 1.7 1
Physalis Solitary rotate AC HM DC Yellow Yellow Yellow A PN F 3.6 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
90
philadelphica Lam. and Black
Plantago lanceolata
L. Spike Irregular ZY HM MC Cream Cream Cream A PL IF 6 3
Plantago major L. Spike Irregular ZY HM MC Cream Cream Cream A PL IF 8 3
Plectranthus
rugosus Benth. Cymose Bilabiate ZY HM MC White White White A PN F 0.7 2
Poa annua L. Panicle Egg ZY HM MC green Cream Cream A PL IF 1 1
Podophylum
haxandrum Royle Solitary Bowl AC HM MC Cream Yellow Yellow A PN F 2.6 2
Polygonum
amphibium L. Spike rotate AC HM MC Pink Pink Pink A PL IF 6 1
Polygonum
amplexicaule Don Spike rotate AC HM MC Pink Pink Pink A PL IF 8 1
Potentilla
argyrophylla Lehm.
Solitary Saucer AC HM MC Yellow Yellow Yellow A PN F 2.5 1
Potentilla reptans L.
Solitary Saucer AC HM MC Yellow Yellow Yellow A PN F 2.4 2
Prunella vulgaris L.
Spike Bilabiate ZY HM DC Purple White White A PN F 0.7 1
Prunus armeniaca
L. Raceme Saucer AC HM MC White Yellow Yellow A PN F 2.2 5
Prunus avium L. Raceme Saucer AC HM MC White Yellow Yellow A PN F 2 3
Prunus domestica L.
Raceme Saucer AC HM MC White Yellow Yellow A PN F 2.4 4
Prunus persica
Batsch Raceme Saucer AC HM MC Pink Red Red A PN F 1.8 4
Pseudomartensia
nemorosa Stewart
and Kazmi
Receme Trumpet AC HM MC Blue yellow yellow A PN F 1.2 1
Chapter – 2 Diversity of floral traits with reference to insect visitors
91
Pteracanthus
urticifolius Bremek.
Spike Campanulate ZY HM MC Purple White White A PN F 1.5 2
Punica granatum L.
Cymose Pear AC HM MC Red Yellow Yellow A PN F 3.2 3
Pyrus communis L. Cymose Saucer AC HM MC White White White A PN F 1.8 3
Pyrus pyrifolia
Nakai Cymose Saucer AC HM MC White White White A PN F 1.6 3
Ranunculus arvensis
L. Cymose Saucer AC HM MC Yellow Yellow Yellow A PN F 1.5 1
Ranunculus laetus
Wall. Cymose Saucer AC HM MC Yellow Yellow Yellow A PN F 2.3 1
Ranunculus
scleratus L. Cymose Saucer AC HM MC Yellow Yellow Yellow A PN F 2.7 2
Raphanus sativus L.
Corymb Bilabiate AC HM MC White Yellow Yellow A PN F 2.4 2
Rhododendron
campanulatum D.
Don
Raceme Campanulate AC HM MC Pink Yellow Yellow A PN F 3.5 2
Robinia
pseudoacacia L Raceme Paplionaceous
ZY HM MC White Yellow Yellow A NC F 1.9 3
Rorippa islandica
Borbas Raceme cruciform AC HM MC Yellow Yellow Yellow A PN F 0.4 2
Rosa brunonii
Lindl. Raceme Rosaceous AC HM MC Pink Yellow Yellow P F 5.5 3
Rosa canina L. Raceme Rosaceous AC HM MC Pink Yellow Yellow P PN F 5.2 3
Rosa indica L. Raceme Rosaceous AC HM MC Red Yellow Yellow P PN F 7 3
Rubus fruticosus L. Raceme Rosaceous AC HM MC White White White A PN F 2 3
Rubus niveus Raceme Rosaceous AC HM MC Pink White White A PN F 1 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
92
Thunb.
Rubus occidentalis
L. Raceme Rosaceous AC HM MC White White White A PN F 1.6 3
Rubus ulmifolius
Schott Raceme Rosaceous AC HM MC Pink Brown Brown A PN F 2.7 3
Rudbeckia hirta L. Capitulum Ligulate and
Tubular Both HM DC Yellow
and Red Yellow Yellow A PN IF 7 4
Rumex acetosa L.
Panicle Rotate AC HM MAC
Yellow
green and
Pink
White White A PL F 0.8 1
Salix alba L. Catkin Catkin AC UN MC Yellow Yellow Yellow A PL IF 7 3
Salix caprea L. Catkin Catkin AC UN MC Yellow Yellow Yellow A PL IF 3 5
Salvia hians Benth. Verticilaster Bilabiate ZY HM MC Blue Yellow Yellow A PN F 2.1 2
Salvia moocroftiana
Benth. Verticilaster Bilabiate ZY HM MC Violet Yellow yellow A PN F 2.6 2
Sambucus wightiana
Wight and Arn. Corymb Rotate AC HM MC White Yellow Yellow A PL IF 12 6
Sanvitalia
procumbens Lam. Capitulum Ligulate and
Tubular Both HM DC Yellow Yellow Yellow A PL IF 2.5 3
Saussurea costus
Lipsch. Capitulum Ligulate and
Tubular Both HM MC Purple Violet cream A PN IF 2.8 3
Senecio
chrysanthemoides
DC.
Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PL IF 2 2
Sisymbrium irio L. Raceme cruciform AC HM MC Yellow Yellow Yellow A PN F 1.3 2
Sium latijugum Umbel Disc AC HM MC White White White A PN IF 8 4
Chapter – 2 Diversity of floral traits with reference to insect visitors
93
Clarke
Skimmia anquetilia
Shaw Panicle Bowl AC HM MC Yellow Yellow Yellow P PN F 0.5 2
Solanum melongena
L. Cymose Rotate AC HM DC Purple Yellow Yellow A PN F 3 2
Solanum nigrum L. Cymose Rotate AC HM DC White Yellow Yellow A PN F 0.7 2
Solanum tuberosum
L. Cymose Rotate AC HM DC White Yellow Yellow A PN F 3 2
Sophora japonica L.
Raceme Paplionaceous
ZY HM MC White Yellow Yellow A PN F 2.1 2
Sonchus oleraceus
L. Capitulum Ligulate ZY HM MC Yellow Yellow Yellow A PL IF 2 1
Stachys floccosa
Benth. Verticilaster Bilabiate ZY HM MC White White White A PN F 1 1
Stellaria media Vill.
Panicle Caryophyllaceo
us AC HM MC White White White A PN F 1 2
Sternbergia lutea
Spreng. Solitary Rotate AC HM MC Yellow Yellow Yellow A PN F 4 4
Syringa emodi
Royle Panicle Hooded ZY HM MC White Yellow Yellow P PL IF 20 6
Tagetes patula L. Capitulum Ligulate and
Tubular Both HM DC Maroon Orange Orange A PL F 3.5 2
Tanacetum vulgare
L. Capitulum Ligulate and
Tubular Both HM MC Yellow Yellow Yellow A PL IF 4 2
Taraxacum
officinale Wigg. Capitulum Ligulate ZY HM MC Yellow Yellow Yellow A PN IF 4 3
Thymus lineasis Verticilaster Bilabiate AC HM MC Purple White White A NC F 0.6 2
Chapter – 2 Diversity of floral traits with reference to insect visitors
94
Benth.
Thymus serphyllum
L. Verticilaster Bilabiate AC HM MC Purple White White A NC F 0.6 2
Trifolium pratense
L.
Pedunculate
head Paplionaceous
ZY HM MC Pink Dark
golden
Dark
golden A PN IF 2.5 4
Trifolium repens L. Pedunculate
head Paplionaceous
ZY HM MC White Dark
golden
Dark
golden A PN IF 2.4 4
Tulipa stellata
Hook. Solitary Campanulate AC HM DC
White and
Red Violet Black A PN F 4.5 4
Verbascum thapsus
L. Spike Rotate AC HM MC Yellow Yellow Yellow A PN F 1.5 1
Veronica arvensis
L. Solitary Rotate AC HM MC Blue White White A PN F 0.8 2
Veronica persica
Poir. Solitary Rotate AC HM MC Blue White White A PN F 0.8 2
Viburnum
grandiflorum Wall.
ex DC.
Corymb Trumpet AC HM DC
White and
Pink Cream cream P PN F 0.8 3
Viola biflora L. Solitary Irregular ZY HM MAC Yellow Yellow Yellow A NC F 3 1
Viola odorata L. Solitary Irregular ZY HM MC Violet Yellow Yellow P NC F 3 1
Weigela floribunda
C.A.Mey. Cymose Tubular AC HM MC Pink White White A PN F 3 2
Zea mays L. Spike Egg ZY UN MC Yellow Yellow Yellow A PL IF 30 17
Zinnia elegans L. Capitulum Ligulate AC HM DC Pink Yellow Yellow A PN IF 8 6
Chapter – 2 Diversity of floral traits with reference to insect visitors
95
DISCUSSION
The results of the present study showed that there is a great diversity in
floral traits including both types of traits (inflorescence as well as flower)
which include inflorescence architecture, inflorescence/flower display size,
flower reward, flower form, flower colour, flower shape and flower symmetry
(Table 2.3; Figs. 2.1-2.12) is in conformity with Waser and Chittka in 1998.
Studied pollen and nectar producing plants showed diversity in floral traits both
attracting as well as rewarding one (Table 2.3, Figs. 2.1-2.12) is in accordance
with Faegri and van der Pijl in 1979. The present study revealed that flowers
act as advertisement units for a variety of pollinators; advertisement efficacy of
flowers varies due to the variation in floral traits (Table 2.3, 4.1) is in alliance
with the study of Raguso (2004). Different sets of floral traits were noted
functioning for recruiting of different types of pollinators accessing some
pollinators and excluding others structuring the dimensions of flower pollinator
web of a region (Table 2.3, 4.1) is in accordance with Faegri and van der Pijl
1979. Flower advertisement policy is the reflection of quality and quantity
floral traits displayed for pollinator recruitment for effective pollen transfer
(Table 2.3, Figs. 2.1-2.12)(Dafni 1992). The quantity and quality of floral
traits affects the pollinator selection, foraging behaviour and its efficacy as
pollen vector at the flower (Stebbins 1970; Dewenter et al., 2005). It is found
(Table 2.3, 4.1) that diversified floral traits react with the diverse pollinator
fauna framing flower-pollinator web with wide range is in conformity with
Armbruster (1984). Research findings revealed a co-relation between floral
traits both attracting and rewarding with pollinator visitation rate (Conner et
al., 1996; Hegland et al., 2005) and reflect a great relationship between floral
traits and pollinators visitation rate (Bosch et al., 1997; Fulton and Hodges,
1999) (Table 2.3). Pollinator uses all types of cues a flower reflects but visual
and olfactory cues are mostly used to address a flower (Raguso and Willis,
Chapter – 2 Diversity of floral traits with reference to insect visitors
96
2002). Floral trait characteristics affect the foraging decision and behaviour of
pollinator at a flower supported by Sargent and Otto 2006.
The present study revealed that there is a great diversity in inflorescence
architecture among the studied plant species and maojority of plant species
displayed raceme followed by cymose, capitulum etc. (Fig. 2.1). It is most
likely that umbel, capitulum and solitary inflorescences with larger bowl-
shaped display has a general advertisement policy and receives maximum
number of flower visits suggesting that greater the inflorescence display size,
greater is the pollinator visitation rate and vice versa (Table 4.1). Inflorescence
architecture is a main driver of pollinator attraction and plays a very special
role in pollinator behaviour at an inflorescence. The larger the inflorescence
size, higher is the pollinator visitation rate as obvious from the observations
that Zea mays has largest functional inflorescence and has the greatest
pollinator visitation rate (Table 2.3) coincides with the study of Thompson in
2001, Harder et al., in 2001, Ohashi and Yahara 2001. Inflorescences are more
attractive as compared to individual pollinators is in accordance with Malerba
and Nattero in 2011. Inflorescence display increase the attractiveness of
pollinators and thus increase visitation rate of pollinators and pollinator
movement varies with different inflorescence architecture is in accordance with
Lawrence et al. (2004). Increase in pollinator visitation rate and pollen export
in large displayed inflorescence revealed the direct relation between
inflorescence display size and pollinator visitation rate is in alliance with Delph
et al., 2004.
The present results revealed that among 215 plant species floral display
was measured for 155 plant species where as inflorescence display was
measured for 60 plant species suggesting that majority of plant species were
displaying flowers as advertisement units. Among different floral display
classes studied, majority of plant species are belonging to small floral display
class (Table 2.1, Fig. 2.2) and among studied inflorescence display classes,
Chapter – 2 Diversity of floral traits with reference to insect visitors
97
majority of plant species are belonging to very small inflorescence display
class (Table 2.2, Fig. 2.3). Among a set of floral traits a flower displays for
pollinator recruitment, floral /inflorescence display size was found as a vital
trait for pollinator recruitment is in agreement with most of studies (Conner et
al., 1996, Willson and Bertin 1979, Grindeland et al., 2005) and corolla display
size was found an important driver for pollinator visitation rates (Bell, 1985)
(Table 2.3). Findings from the observation revealed the flower/inflorescence
display size plays very important role in decision making of pollinator for
visitating a flower is in accordance with Medel et al. (2003). Flower display
and flower symmetry are the important floral attributes which affects the
pollinator visitation rate: flower symmetry of studied plant species varied,
majority were actinomorphic and generalized, and a lesser percentage was
zygomorphic and specialized which architectures their pollinator fauna (Table
2.3 and Table 4.1; Fig. 2.6) is in accordance with the study of Hegland and
Totland in 2005).
Results revealed that reward presentation by studied plant species are of
three types (i) pollen presentation (ii) nectar presentation (iii) both and among
studied pollinator flora majority of plant species are presenting both types of
rewards (Fig. 2.4). It is evident from the results that every pollinator is guided
and signalled by the floral traits towards the ultimate reward that is in the form
of pollen and nectar is in accordance with Waser in 1983. Enlisted plant species
diverging in rewards, prevailing pollen and nectar as a common reward (Fig.
2.4) can serve for commercial bee- keeping industry at large scale supported by
Dukku 2013. Pollen presentation with attracting colour acts as a strong signal
for attraction of pollinators towards the flower and modifies the decision and
foraging behaviour of a pollinator at a flower as present result revealed that
Salix caprea are more attractive and has more pollinator visitation rate (PVR)
than S. alba, and both species differs in their advertisement efficiencies (Table
2.3), such type of studied was also carried by Dotterl and Vereecken 2010. A
Chapter – 2 Diversity of floral traits with reference to insect visitors
98
great diversity in floral shape was noted among the studied plant species (Fig.
2.5) is in accordance with Waser and Chittka in 1998. Among diverse floral
shapes observed, majority of pollinator flora displaying rotate flower shape
(Fig. 2.5) but according to flower symmetry, flower shapes are of two types
open access shapes (actinomorphic) and closed access shapes (zygomorphic)
and it was found that majority of plant species had open access flower shapes
such as bowl, saucer reflecting general advertisement policy and thus received
greater number of pollinator taxa (Fig. 2.6, Table 4.1). Closed access flowers
like trumpet, paplionaceous had specialized advertisement policy and received
less number of pollinator taxa, that restrict the access to general categories but
provided access to specialized pollinators only, thus avoiding the nectar
robbers or cheaters by excluding non efficient pollinators with the help of
specialized floral traits (Table 2.3 and Table 4.1) is in conformity with the
study of Raguso (2004) and Guerrero et al. (2014). Flowers with specialized
traits like asymmetrical and zygomorphic corolla tube select the specialized
type of pollinator for pollen export thus reflecting the pollination syndrome.
The present results revealed that among 215 plant species majority were
bisexual suggesting that they serve as general, potential and wide source of
reward for both pollenivorous as well as nectarivorous flower visitors as
compared to unisexual which are narrow in rewrad range (Fig. 2.7). Male
unisexual plant species were found to receive a higher number of pollinator
visits because presence of high level of reward including both pollen and
nectar. Castanea sativa (07), Salix caprea (05), Zea mays (05), Actinidia
deliciosa (04). Carya illinoinensis (03), Ailanthus altissima (02) and Salix alba
(02) (Fig. 2.4). It was found that female unisexual flowers produce only nectar
as a reward. Bisexual and male unisexual flowers produce pollen and nectar as
a reward for pollen vector. Sex of pollen and nectar producing plants revealed
that the percentage of uni-sexuality in plant species acting as forage plants is
rare and it is found that out of 215 forage plants, 208 (97%) are bisexual while
Chapter – 2 Diversity of floral traits with reference to insect visitors
99
as only 07 plant species (3%) are unisexual shows alliance (Table 2.3, Fig. 2.7)
with the studies of Richard (1997) and Renner and Ricklefs (1995), showed
only 6 % of flowering plants have separate male and female flowers and this
phenomena is called dioecy. Male flowers are investing more for pollinator
attraction as compared female flowers (Ashman, 2009). It was an important
observation that after the flower anthesis, flower visitors prefer nectar as
compared to pollen. Present study finding revealed that the unisexual plants
had different sex on different plants with different rewards composition.
Reward presentation of flower varies between male flowers and female
flowers. Male unisexual plants produced both pollen and nectar as a reward but
female unisexual flowers produce nectar only as a reward (Table 2.3) is in
accordance with the study of Vaughton and Ramsey 1998. Nectar is the only
floral reward which both male and female flowers present for pollinator
attraction (Table 2.3). It is found that generally female flowers has less
visitation rate as compared to male flowers due to differential potential in
reward presentation (Delph and Ashman, 2006; Blair and Wolfe, 2007).
Advertisement is the important cue for both male and female flowers for
attracting pollinators (Raguso, 2008).
There was not a significant relation found between flower chromatism
and visitation rate though other floral traits such as flower display size affects
the pollinator visitation rate (Fig. 2.8). Among studied pollinator flora majority
showed multichromatism of their flowers (Fig. 2.8). A large diversity in flower
colour is displayed by the studied pollen and nectar producing plants, among
which majority of plant species displaying yellow colour flowers (Fig. 2.9) and
pollinator visitation rate for yellow coloured flowers was found greater than
other flower colours (Table 2.3 and 4.1) is supported by Campbell et al., 2010.
Flower colour a main cue for attraction of pollinators includes colour of corolla
tube, anther colour and pollen colour as pollen loaded flowers are perceived
more rapidly, thus pollen colour acts as a strong cue to affect the pollinator
Chapter – 2 Diversity of floral traits with reference to insect visitors
100
behaviour at the flower is in conformity with the study of Lepage and Boch
(1968).
CONCLUSIONS
The present study revealed that there is a great diversity within the floral
traits a flower displays for the attraction of pollinators. Attracting floral traits
guide and direct the flower visitor towards the ultimate reward which serves as
energy source for pollinators. Floral traits include both traits of an
inflorescence and its flowers. Study of inflorescence architecture of pollinator
flora showed a great diversity with majority of plant species with raceme
inflorescence. Floral display size is the most important driver of pollinator
attraction, showed a great diversity categorized into 5 classes each with
specific range of display size such as very small floral display (VSFD), small
floral display (SFD), medium floral display (MFD), large floral display (LFD),
very large floral display (VLFD) and great relation between floral display size
and pollinator visitation rate was observed, greater the display size greater the
pollinator visitation rate and vice versa. Infloresecnec display wasalso five
categories such as VSID, SID, MID, LID and VLID and among these majority
of pollinator flora has VSID. As compared to single flower, inflorescence
attracts the more number of flower visitors due to increased inflorescence
display size. Majority of pollen and nectar producing plant species were
bisexual serving source for both pollen and nectar.
Flower shape is an important floral trait which is a main factor in
determining the foraging behaviour. A great diversity in flower shape was
observed and majority of pollinator flora has rotate shaped flowers. Analysis of
flower symmetry of pollinator flora revealed that majority of plants has
actinomorphic flowers suggesting generalized approach towards pollinators.
Generally two flower shapes were observed open access shapes include rotate,
saucer, bowl, cruciform, rosaceous, and closed access flower includes bilabiate,
Chapter – 2 Diversity of floral traits with reference to insect visitors
101
campanulate, trumpet, paplionaceous and caryphylaceous, personate. These
two shaped flowers have different interacting efficacies with pollinators and
thus modify generalization-specialization continuum range of interaction.
Flower symmetry concludes that symmetry flowers act as generalized source
and has more pollinator visitation rate as compared to asymmetrical flowers.
Majority of flowers were found to show monochromatism as a floral trait
followed by dichromatism and multichromatism. Flower colour too had a great
diversity and majority of pollinator flora has yellow coloured flowers. Anther
and pollen colour were also found to show diversity among documented pollen
and nectar producing plants and majority of pollinator flora displays yellow
colured anthers and pollens respectifully. Floral scent presentation among
studied pollinator flora revealed that majority of plant species were without
scent.
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INTRODUCTION
Usually flower advertisement attracts an assemblage of insect visitors.
Among these insect visitors, some are more efficient pollen vectors
(pollinators), others are inefficient and few are robbers which include
butterflies and moths; the latter rob the flower reward without aiding in pollen
transfer (Thompson, 1982; Bronstein, 1994). Nectar robbers are described as
cheaters in plant pollinator mutualisms due to their cheating habit of extracting
floral rewards without providing service.
Owing to global decline in pollinators, management of pollinators for a
wide range of crops generates need to search for alternative, efficient, native
pollinator for sustainable pollination. So the preliminary task is to test the
validity of alternative pollinator for a particular plant species and avoid
concentrating on all the flower visitors, as pollinators vary in their pollen
deposition capacities on receptive stigma and visitation rate. One of the well-
known methods in pollination biology is to test the pollinator effectiveness by
comparing principle pollinators from those who just are flower visitors or
flower cheater/robbers (Popic et al., 2013). Ecological and evolutionary
prospective of pollination network has made its analyses a current challenge for
its sustainability with the climate change (Willmer, 2012).
For categorizing flower visitors into efficient and inefficient pollinators,
analysis of pollen vector efficacy by single visit pollen deposition (SVD) on
virgin stigma is considered to be the most validated approach in addition to
visitation rate (King et al., 2013). The method reveals the potential pollinators
for different foraging plant species and also has application in guiding towards
the conservation and sustainable use in pollinator deficit ecosystems. The SVD
method avoids the measurement of fruit and seed set during post-pollination
and provides the accurate pollen carrying potential of pollinators. In recent
times, pollinator decline as well as deteriorating quality and quantity of
pollinator dependent ecosystems as generated interest in studying pollinator
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diversity, occurrence and effectiveness. The pollinator effectiveness (PE) is the
pollen carrying and depositing capacity of pollen vectors (Neeman et al.,
2010). Pollinator ecology in terms of pollinator effectiveness has been studied
in recent years at community level in order to understand the pollinator deficits
and its affect on interacting population (Waser et al., 1996), as pollinator
deficit is the main cause of deterioting quality as well as quantity in
ecosystemic outcome. Therefore, the present study makes an attempt to to
categorise efficient and inefficient pollinators.
REVIEW OF LITERATURE
Research on pollinator effectiveness has been carried throughout the
world in order to describe principle pollinators for different crops and different
pollinators were found to have different pollen carrying potential for different
plant species (Free, 1970). Torchio (1990) found that evaluation of flower
visitor’s role enables in making informed pollinator choice for maximization of
crop pollination. Despite the importance of managed agricultural pollinators as
pollen vectors, the pollen carrying and depositing potential of different natural
pollinators is well established. Thus, it has been advised to search for
alternative, less vulnerable, indigenous and compatible pollinator for sustaining
pollination, as over dependence on managed pollinator, such as Apis mellifera
for wide range of crop pollination is unsustainable due to the global decline in
its population (Tepedino, 1981; Torchio, 1990).
Stephen (1962) and Bohart (1972) studied pollination effectiveness of
Megachile rotundata and found that it acts as an effective pollen vector for
Medicago sativa. Ruijter (1997) studied the pollen carrying and depositing
ability of bumble bees on Lycopersicon esculentum and found that they
function as efficient pollinator for Lycopersicon esculentum. Thomson and
Goodell (2001) studied the effectiveness of pollinators (Apis and Bombus) on
Chapter – 3 Determination of pollinator effectiveness among insect visitors
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apple and almond and found that bumblebee visitors were found to deposit
more pollen on virgin stigmas of apple and almond as compared to honeybee
visitors. Castellanos et al. (2003) studied pollen deposition by bumblebees and
hummingbirds on bird pollinated Penstemon barbatus plant and bee pollinated
P. strictus. They found that bees and birds have similar pollen transfer
efficacies and bird pollinated plant species had more pollen deposited as
compared to bee pollinated plants because pollen on bee bodies become
quickly unavailable for transfer.
Pollinator effectiveness among different flower visitors and their
comparison for categorizing them into different groups were made by different
workers (Primark and Silander, 1975; Herrera, 1987; Waser and Price, 1990).
Faegri and Van der Pijl (1979) found that floral trait sets which are traditionally
associated with pollinators, guides and predicts the pollinators for a particular
set of floral traits and makes pollinator a best pollen vector for a particular
plant. Thomson (2000) found that different pollination syndromes predict
different pollinators due to their different pollinator effectiveness or pollen
carrying capacity.
Wolfe et al. (1989) studied pollen deposition on the stigma of
Pontederia cordata and found that different rates of pollen deposition with
respect to variety of plant species. Freitas and Paxton (1998) studied
comparison of pollen carrying and depositing capacity between exotic Apis
mellifera and indigenous Centris tarsata on Anacardium occidentale and found
that Centris tarsata is more efficient pollen depositor as compared to Apis
mellifera: pollenivorous visitor of any species were more efficient in
extracting, caring and depositing pollens as compared to nectarivorous member
of the same species.
Young and Stanton (1990) found that pollen deposition on stigma is one
of the important factors for achieving more reproductive success in plants.
Burkle et al. (2007) found that among the different categories of flower
Chapter – 3 Determination of pollinator effectiveness among insect visitors
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visitors, reward robbers especially nectar robbers have negative effects on plant
reproductive success by robbery only nectar without aiding in pollen transfer.
Spira et al. (1992) studied the pollen deposition on the receptive stigma by
SVD method on Hibiscus spp. and found that median number of pollen grains
deposited on the stigma per visit was seventy pollen grains and the flowers
were recorded to have four visits per hour. Bingham and Orthner (1998)
studied pollination mechanism in alpine and foothill populations of Campanula
rotundifolia by observing different objectives like visitation rate, pollen
deposition on stigma and stigma receptivity.
Corner et al. (1995) studied pollination effectiveness among four
different flower visitor taxa and found that they vary in pollen carrying and
depositing capacity by investigating single visit pollen deposition on stigma.
Caban and Ackerman (2007) studied behaviour of flower vistors and their
pollination effectiveness and found that among different flower visitors long
billed hummingbird was efficient pollinator in terms of pollen deposition.
Wilson and Thomson (1991) studied pollination effectiveness with foraging
behaviour of honeybees and bumblebees on Impatiens capensis and found that
bumblebees were efficient flower visitors with high rates of pollen deposition
as compared to honeybee. Young et al. (2007) studied the relationship between
foraging behaviour and pollen deposition capacity of honey bees and
bumblebees on Impatiens capensis and found that foraging behaviour is one of
the important factors in predicting the pollinator effectiveness of a flower
visitor. King et al. (2013) studied the pollinator effectiveness for 13 temperate
and tropical plant species by single visit pollen deposition method and found
that it was in accordance with pollination syndrome which is based on floral
advertisement and reward.
Whelan et al. (2009) studied foraging behaviour, pollen extracting and
depositing capacity of honeybees and found that honeybees were frequent
visitors and extracted more pollens than deposited at the virgin stigma.
Chapter – 3 Determination of pollinator effectiveness among insect visitors
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Honeybees show a great variation in the efficiency of pollination (Celebrezze
and Paton, 2004), even though collecting both pollen and nectar from different
plant species yet they are reward robbers for many vertebrate plant species
(Gross and Mackay, 1998). Balam et al. (2012) studied the contribution of
honeybees, flies and wasps to pollination of Persea americana and found that
flies carried much more pollen than honeybees and wasps and number of pollen
grains deposited on stigma by each taxa was same. Willmer and Finlayson
(2014) studied the pollination effectiveness of Bombus terrestris workers on
Vinca minor flowers by single visit deposition on virgin stigmas and found that
the pollen deposition is positively related with the bee’s body size, larger bees
deposited more pollen and smaller bees deposited less pollen.
Pollinator effectiveness of any flower visitor is the measure of quality
as well quantity of pollinator’s pollen caring and depositing capacity (Olsen,
1997) and sometimes pollinator effectiveness can be best for most common
pollinators (Ivey et al., 2003) and sometimes pollinator effectiveness can be
best for uncommon pollinators rather than common one (Mayfield et al., 2001).
Generally robbers are not pollinators (Inouye, 1980): in fact many robber
species appear to be pollinators (Guitian et al., 1993). Maloof and Inouye
(2000) showed negative relation between robbers and plant reproductive
success, but several studies in last 5 decades revealed that the effect of robbers
is complex and depend on robber identity, legitimate pollinator identity, and
quantity of flower resource robbery. Nectar robbery occurs due to the
morphological mismatch between robbers and their host flowers (Inouye,
1983). Plant pollinator interaction is found to be susceptible to nectar robbers
due to their cheating role in this particular interaction by taking reward without
pollen transfer (Boucher et al., 1982). Pollen deposition by carpenter bee on
mismatched bird pollinated plant species have been studied and it was found
that robbers are not cheaters but they help in pollen transfer a little (Waser,
1979; Scott, 1989) . Heinrich and Haven (1972) studied the influence of robber
Chapter – 3 Determination of pollinator effectiveness among insect visitors
116
on the behaviour of legitimate pollinator and found that plant reproductive
success is influenced with robbers by changing the behaviour of legitimate
pollinators. Kendall and Smith (1976) studied the validity of robbers on
Phaseolus vulgaris and found that robbers are not entirely cheating flowers
instead caring and depositing a small quantity of pollen grains. Goulson et al.
(1998) studied the behaviour of bumblebees towards robbed and non robbed
flowers and found that robbers influence the behaviour of pollinators because
pollinators are capable to distinguish rewarding (non-robbed) and non
rewarding (robbed) flowers of the same species.
Snow and Roubik (1987) studied the pollinator effectiveness of two
species of genus Centris and found that Centris longimana deposited more
pollen grains on stigma of Cassia flower than Centris inermis. Jones et al.,
(1966) studied the efficacy of two species of bumblebees (long tongued and
short tongued) on Trifoliun pretense and found that long tongued bumblebee
was recognised as legitimate pollinator and short tongued as robber. The
observation from Jones et al. (1966) revealed short tongued bumblebee as not
robbers but less efficient pollinators. Gass and Montgomerie (1981) studied the
behaviour of hummingbird at a flower and found that they are able in
determining the nectar level of a flower and can avoid robbed or empty flowers
as compared to lightly robbed one (Irwin and Brody 1998).
Laverty (1994) studied behaviour of bumble bees and found that the
learning of foraging behaviour for new flower species involves investment of
energy and time of bumblebees that avoid switching to them and favours
foraging of already learned flowers even though they are robbed, enhancing the
plant fitness of learned flowers. Stein and Hansen (2011) studied the behaviour
and ecology of flower visitors of Heliconia angusta and found that it is an
ornithophilous syndromed plant species with Trigona sp., four species of
hummingbirds and two species of butterflies as flower visitors, among them
Chapter – 3 Determination of pollinator effectiveness among insect visitors
117
hummingbirds act as principle pollinators and Trigona sp., butterflies as
robbers of pollen respectively.
MATERIALS AND METHODS
During the present study, both field observations and laboratory
methods were employed in different months of the year 2014-2015 in the study
area. For the field observations, visit centred method was used (Stebbins 1970,
1974; Armbruster, 1988; Armbruster et al., 2000) which quantifies the number
of pollinator visits for a particular flower or inflorescence per 10 minutes. The
event of flower-visitor interaction was photographed (Sony DSC-S2000) for
further processing and study in the laboratory. The field observations were
supplemented with the lab-based methods which involves quantification of
pollens on the virgin stigmas. Pollen carrying potential of different pollinators
was assessed by single visit pollen deposition (SVD) technique on the virgin
stigma by pollinators (Inouye, 1994). The virgin stigma was ensured by
emasculating the flower bud in order to eliminate self-pollination and favour
the foreign pollen deposition with the help of pollen vector. The emasculated
flower buds were caged by bags in order to exclude flower visitors. Flower
buds were selected randomly. The half- opened flower was debagged and the
first flower visitor foraging a particular flower was collected and identified.
The flowers were taken carefully to laboratory and the pollens on stigamas
were observed under the stereozoom microscope (Nikon SMZ 800) and
photographed. Then the pollen grains deposited on the virgin stigma by the
particular flower visitor were counted (Inouye, 1994; Freitas and Paxton, 1998;
Castellanos et al., 2003; Thomson and Goodell, 2011; King et al., 2013;
Primark and Silander, 1975; Herrera, 1987; Waser and Price, 1990; Neeman,
2010).
Chapter – 3 Determination of pollinator effectiveness among insect visitors
118
The present study was conducted using eight (8) different types of insect
visitors, namely: Xylocopa valga, Apis mellifera, Apis cerana, Bombus
tunicatus., Eristalis tenax, Andrena flavipes, Lasioglossum margenatum and
Vanessa cashmeriansis; on 25 flowering plant species which were visited by
these 8 insect visistors, which include: Abelia grandiflora, Actinidia deliciosa,
Aesculus indica, Alcea rosea, Antirrhinum majus, Berberis lycium, Brassica
compestris, B. rapa, Cichorium intybus, Colchicum luteum, Convolvulus
arvense, Coriandrum sativum, Cydonia oblonga, Epilobium hirsutum,
Hypericum hookerianum, H. perforatum, Lavatera cashmiriana, Mahonia
borealis, Malus domestica, Prunus armeniaca, Punica granatum, Rosa
webbiana, R. indica, Sisymbrium irio and Sternbergia vernalis (Table 3.1 and
3.2).
RESULTS
In the present study, pollen depositing ability and visitation rate of eight
different insect visitor species were determined on 25 different foraging plant
species and it was found that all the eight insect visitor species vary in their
pollen depositing as well as visitation rate (Table 3.1, 3.2 and photoplates 3.1,
3.2).
Chapter – 3 Determination of pollinator effectiveness among insect visitors
119
It was found that five different flower visitor species namely Xylocopa
valga, Bombus tunicatus, Apis mellifera, Apis cerana and Vanessa
cashmeriansis were foraging on Abelia grandiflora flowers. By using SVD
technique, it was found that Xylocopa valga deposits highest number (24) of
pollens on the virgin stigma of Abelia grandiflora followed by Bombus
tunicatus (16), Apis mellifera (13), Apis cerana (07) and none by Vanessa
cashmeriansis. Visitation rate was found highest for Apis mellifera (04
visits/10 minutes) followed by Xylocopa valga (03), Bombus tunicatus and
Apis cerana (02 each) and once by a Vanessa cashmeriansis (Fig. 3.1).
Figure 3.1: Pollen deposition and visitation rate of different flower visitors
on Abelia grandiflora.
0
5
10
15
20
25
30
Xylocopa valga Bombus
tunicatus
Apis mellifera Apis cerana Vanessa
cashmeriansis
Pollens deposited
Visits
Number
Pollinator species
Chapter – 3 Determination of pollinator effectiveness among insect visitors
120
Two different flower visitor species namely Apis mellifera and Apis
cerana were found foraging on Actinidia deliciosa. Apis mellifera was found
depositing (28) pollens where as Apis cerana (21) on virgin stigmas of
Actinidia deliciosa and highest visitation rate was observed for Apis mellifera
(12) and (08) for Apis cerana (Fig. 3.2)
Figure 3.2: Pollen deposition and visitation rate of different flower visitors on
Actinidia deliciosa.
Two species namely Apis mellifera and A. cerana were found foraging
on Aesculus indica and were found depositing (07) and (03) pollens on virgin
stigmas of Aesculus indica respectively with highest visitation rate (06) in Apis
cerana and (05) in Apis mellifera (Fig. 3.3).
Figure 3.3: Pollen deposition and visitation rate of different flower visitors
on Aesculus indica.
0
5
10
15
20
25
30
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
1
2
3
4
5
6
7
8
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
121
Apis mellifera and A. cerana were found foraging on Alcea rosea
depositing (13) and (11) pollens respectively with highest visitation rate (06) in
Apis mellifera followed by (03) in Apis cerana (Fig. 3.4).
Figure 3.4: Pollen deposition and visitation rate of different flower visitors on
Alcea rosea.
On the flowers of Antirrhinum majus, two different flower visitor
species were found foraging. Xylocopa valga were found depositing highest
(16) pollens followed by Apis mellifera (07) on the virgin stigmas of A. majus.
Xylocopa valga had high (05) visitation rate than A. mellifera (03) (Fig. 3.5).
Figure 3.5: Pollen deposition and visitation rate of different flower visitors on
Antirrhinum majus
0
2
4
6
8
10
12
14
Apis mellifera Apis cerana
Pollens deposited Visits
Number
Pollinator species
0
2
4
6
8
10
12
14
16
18
Xylocopa valga Apis mellifera
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
122
Pollen depositing efficacies of three different taxa was tested on stigmas
of Berberis lycium by SVD technique and it was found that Apis mellifera
deposited highest (23) number of pollens followed by Apis cerana 18 and
Eristalis tenax (4) on virgin stigmas of Berberis lycium. Highest visitation rate
was recorded for Apis cerana (4) followed by rest other three taxa having (03)
each (Fig. 3.6).
Figure 3.6: Pollen deposition and visitation rate of different flower visitors on
Berberis lycium.
The pollinating abilities of five different flower visitor species were
investigated on Brassica compestris by SVD and found that Apis mellifera
deposited highest (15) number of pollens followed by Apis cerana (11),
Andrena flavipes (8), Eristalis tenax (5) and Lasioglossum margenatum (4).
Lasioglossum margenatum has highest visitation rate (06) followed by Apis
cerana (04), Apis mellifera (03), Andrena flavipes (02) and (01) of Eristalis
tenax (Fig. 3.7).
Figure 3.7: Pollen deposition and visitation rate of different flower visitors on
Brassica compestris.
0
5
10
15
20
25
Apis mellifera Apis cerana Eristalis tenax
Pollens deposited Visits
Pollinator species
Number
0
2
4
6
8
10
12
14
16
Apis
mellifera
Apis cerana Andrena
flavipes
Eristalis
tenax
Lasioglossum
margenatum
Pollens deposited Visits
Numbe
r
Pollinator species
Chapter – 3 Determination of pollinator effectiveness among insect visitors
123
The SVD analysis revealed that Apis mellifera deposited highest (15)
number of pollens followed by Apis cerana (9), Andrena flavipes (5) and
Lasioglossum margenatum (3) on the virgin stigmas of Brassica rapa.
Visitation rate (5 visits/10 minutes) was recorded highest for Lasioglossum
margenatum and Apis cerana followed by Apis mellifera (4) and (03) Andrena
flavipes (Fig. 3.8).
Figure 3.8: Pollen deposition and visitation rate of different flower visitors on
Brassica rapa.
Pollen deposition analysis revealed that among three different flower
visitor species, Apis mellifera deposited highest number of pollens (20)
followed by Apis cerana (13) and (06) by Eristalis tenax on the virgin stigmas
of Cichorium intybus. Visitation rate for Apis mellifera (02) was found more
than rest three having (01each) (Fig. 3.9).
Figure 3.9: Pollen deposition and visitation rate of different flower visitors on
Cichorium intybus.
0
2
4
6
8
10
12
14
16
Apis mellifera Apis cerana Andrena flavipes Lasioglossum
margenatum
Pollens deposited Visits
Number
Pollinator species
0
5
10
15
20
25
Apis mellifera Apis cerana Eristalis tenax
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
124
On the flowers of Colchicum luteum, pollen deposition and visitation
rate by two species of Apis was observed and it was found that Apis mellifera
deposited more pollen (18) as compared to Apis cerana (12). Visitation rate
was observed highest for Apis cerana (06) and (05) for Apis mellifera (Fig.
3.10).
Figure 3.10: Pollen deposition and visitation rate of different flower visitors on
Colchicum luteum.
Pollen deposition on virgin stigmas of Convolvulus arvense revealed
that Apis mellifera deposited highest (21) number of pollens as compared to
Apis cerana deposited (16) and visitation rate revealed that Apis mellifera (02)
visits twice the rate of (01) Apis cerana (Fig. 3.11).
Figure 3.11: Pollen deposition and visitation rate of different flower visitors
on Convolvulus arvense.
0
5
10
15
20
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
5
10
15
20
25
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
125
Pollen deposition on virgin stigmas of Coriandrum sativum by three
different flower visitor species revealed that Apis mellifera depositing highest
number of pollens (12) followed by Apis cerana (08), Andrena flavipes (05).
Visitation rate revealed that Apis cerana and Andrena flavipes has highest rate
(05 each) followed by Apis mellifera (04) (Fig. 3.12).
Figure 3.12: Pollen deposition and visitation rate of different flower visitors on
Coriandrum sativum.
Results of pollinator effectiveness on the Cydonia oblonga flowers
revealed that Apis mellifera deposited (11) pollens with visitation rate (03) and
Apis cerana deposited (09) pollens with visitation rate (02) on virgin stigmas of
Cydonia oblonga (Fig. 3.13).
Figure 3.13: Pollen deposition and visitation rate of different flower visitors
on Cydonia oblonga.
0
2
4
6
8
10
12
14
Apis mellifera Apis cerana Andrena flavipes
Pollens deposited Visits
Pollinator species
Number
0
2
4
6
8
10
12
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
126
Apis mellifera was found to deposit (31) pollens and Apis cerana
deposited (24) on the virgin stigmas of Epilobium hirsutum. Visitation rate
revealed that Apis mellifera has visitation rate twice the rate of Apis cerana
(Fig. 3.14).
Figure 3.14: Pollen deposition and visitation rate of different flower visitors
on Epilobium hirsutum.
Pollen deposition on the flowers of Hypericum hookerianum and H.
perforatum by two species of Apis revealed that Apis mellifera deposited (23)
and (22) pollens whereas Apis cerana deposited (18) and (16) pollens on their
virgin stigmas respectively. Visitation rate revealed that Apis mellifera showed
highest rate (05) and (04) for H. hookerianum and H. perforatum respectively
whereas Apis cerana showed (02) for each (Figs. 3.15, 3.16).
Figure 3.15: Pollen deposition and visitation rate of different flower visitors
on Hypericum hookerianum.
0
5
10
15
20
25
30
35
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
5
10
15
20
25
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
127
Figure 3.16: Pollen deposition and visitation rate of different flower visitors
on Hypericum perforatum.
Pollen deposition in virgin stigmas of Lavatera cashmiriana revealed
that Apis mellifera deposited highest (09) number of pollens and Apis cerana
deposited (07). Visitation rate revealed that Apis mellifera has visitation rate
(05) as compared to Apis cerana which had (02) (Fig. 3.17).
Figure 3.17: Pollen deposition and visitation rate of different flower visitors
on Lavatera cashemiriana.
0
5
10
15
20
25
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
1
2
3
4
5
6
7
8
9
10
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
128
Pollen deposition on Mahonia borealis revealed that Apis mellifera
deposited highest (28) number of pollens followed by Apis cerana (16).
Visitation rate revealed that both had equal (03) visitation rates (Fig. 3.18)
Figure 3.18: Pollen deposition and visitation rate of different flower visitors on
Mahonia borealis.
Pollen deposition on virgin stigmas of Malus domestica by five different
flower visitors revealed that Xylocopa valga deposited (23) highest number of
pollens followed by Apis mellifera (17), Apis cerana (12), Andrena flavipes
(08) and Vanessa cashmeriansis was not found to deposit any pollen. Visitation
rate revealed that Apis mellifera has highest visitation rate (04) followed by
Apis cerana (03), Xylocopa valga, Andrena flavipes (02) each and finally by
Vanessa cashmeriansis once (Fig. 3.19)
Figure 3.19: Pollen deposition and visitation rate of different flower visitors on
Malus domestica.
0
5
10
15
20
25
30
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
5
10
15
20
25
Xylocopa valga Apis mellifera Apis cerana Andrena
flavipes
Eristalis tenax Vanessa
cashmeriansis
Pollens deposited Visits
Number
Pollinator species
Chapter – 3 Determination of pollinator effectiveness among insect visitors
129
Pollen deposition on virgin stigmas of Prunus armeniaca by four
different flower visitor species revealed that Xylocopa valga deposited highest
number of pollens (27) followed by Apis mellifera (19), Apis cerana (15) and it
was found that Vanessa cashmeriansis does not deposit any pollen grain.
Visitation rate revealed that Apis cerana has highest visitation rate (08)
followed by Apis mellifera (07) and (03) by Xylocopa valga and finally by
Vanessa cashmeriansis twice (Fig. 3.20).
Figure 3.20: Pollen deposition and visitation rate of different flower visitors on
Prunus armeniaca.
Pollen deposition on virgin stigmas of Punica granatum revealed that
among two different flower visitor species, Apis mellifera deposited more
pollens (39) as compared to Apis cerana (28) and has more visitation rate (3) as
compared to later (02) (Fig. 3.21).
Figure 3.21: Pollen deposition and visitation rate of different flower visitors on
Punica granatum.
0
5
10
15
20
25
30
Xylocopa valga Apis mellifera Apis cerana Vanessa
cashmeriasis
Pollens deposited
Visits
Number
Pollinator species
0
5
10
15
20
25
30
35
40
45
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
130
Pollen deposition on virgin stigmas of Rosa webbiana by three different
flower visitor species revealed that Xylocopa valga deposited highest number
of pollens (23) followed by Apis mellifera (18) and Apis cerana (13). Visitation
rate revealed that both members of Apis has same visitation rate (06) and (02)
for Xylocopa valga (Fig. 3.22).
Figure 3.22: Pollen deposition and visitation rate of different flower visitors on
Rosa webbiana.
Analysis of pollen depositing abilities of Apis mellifera and Apis cerana
revealed that former deposited more pollen as compared to later on the virgin
stigmas of Rosa indica. Visitation rate revealed that Apis mellifera has high
(04) visitation rate than Apis cerana (03) (Fig. 3.23)
Figure 3.23: Pollen deposition and visitation rate of different flower visitors on
Rosa indica.
0
5
10
15
20
25
Xylocopa valga Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
0
2
4
6
8
10
12
14
16
18
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
131
Pollen deposition on virgin stigmas of Sisymbrium irio revealed that
Apis mellifera (18) deposited more pollen than Apis cerana (13). Visitation rate
revealed that Apis mellifera has high visitation rate (02) than Apis cerana (01)
shown (Fig. 3.24)
Figure 3.24: Pollen deposition and visitation rate of different flower visitors on
Sisymbrium irio.
Pollen deposition on the stigmas of Sternbergia vernalis revealed that
Apis mellifera deposited (17) pollens and Apis cerana (13). Visitation rate
revealed that Apis mellifera forages (04) times where as Apis cerana (03) times
(Fig. 3.25).
Figure 3.25: Pollen deposition and visitation rate of different flower visitors
on Sternbergia vernalis.
0
2
4
6
8
10
12
14
16
18
20
Apis mellifera Apis cerana
Pllens deposited Visits
Pollinator species
Number
0
2
4
6
8
10
12
14
16
18
Apis mellifera Apis cerana
Pollens deposited Visits
Pollinator species
Number
Chapter – 3 Determination of pollinator effectiveness among insect visitors
132
A comparative analysis of pollen deposition by Apis mellifera and Apis
cerana on different flowering plants revealed that Apis mellifera deposited
highest number of pollens than A. cerana on all plant species as shown Figure
3.26.
Figure 3.26: Pollen depositions by Apis mellifera and A. cerana on different
plants.
0
5
10
15
20
25
30
35
40
45
Apis mellifera
Apis cerana
No. of pollens deposited
Plant species
Chapter – 3 Determination of pollinator effectiveness among insect visitors
133
Comparative analysis of visitation rate of Apis mellifera and A. cerana
on different flowering plants revealed that Apis mellifera shows highest
visitation rate than A. cerana for 15 different plant species while as Apis cerana
has highest visitation rate for 7 plant species and both displaying equal rates for
2 plant species (Fig. 3.27).
Figure 3.27: Visitation rate of Apis mellifera and A. cerana on different
flowering plants.
0
2
4
6
8
10
12
14
Apis mellifera
Apis cerana
Plant species
No. of viisits
Chapter – 3 Determination of pollinator effectiveness among insect visitors
134
Comparative study of pollen deposition on virgin stigmas and visitation
rates of Xylocopa valga, Apis mellifera and A. cerana on different flowering
revealed that Xylocopa valga deposited the highest number of pollens on virgin
stigmas followed by Apis mellifera and A. cerana and visitation rate varies;
Xylocopa valga shows highest visitation rate for Prunus armeniaca and Apis
mellifera shows highest rate for Abelia grandiflora and Malus domestica (Figs.
3. 28 and 3.29).
Figure 3.28: Pollen depositions by Xylocopa valga, Apis mellifera and A.
cerana on different flowering plants.
Figure 3.29: Visitation rate of Xylocopa valga on different flowering plants.
0
1
2
3
4
5
6
7
8
9
Prunus
armeniaca
Abelia
grandiflora
Malus domestica Rosa webbiana
Apis cerana
Apis mellifera
Xylocopa vaga
No. of visits
Plant species
0
1
2
3
4
5
6
7
8
9
Prunus armeniaca Abelia grandiflora Malus domestica Rosa webbiana
Apis cerana
Apis mellifera
Xylocopa vaga
No. of visits
Plant species
Chapter – 3 Determination of pollinator effectiveness among insect visitors
135
Table 3.1: Pollen deposition on different plant species by different insect visitors
Plant species Family Apis
mellifera
Apis
cerana
Xylocopa
valga
Bombus
tunicatus
Eristalis
tenax
Andrena
flavipes
Lasioglossum
margenatum
Vanessa
cashmeriansis
Abelia grandiflora Rehd. Caprifolaceae 13 7 24 16 - - - 0
Actinidia deliciosa Liang and Ferguson. Actinidiaceae 28 21 - - - - -
Aesculus indica Hook. Sapindaceae 7 3 - - - - -
Alcea rosea L. Malvaceae 13 11 - - - - -
Antirrhinum majus L. Scrophulariaceae
7 - 16 - - - -
Berberis lycium Royle Berberidaceae 23 18 - - 4 - -
Brassica compestris Brassicaceae 15 11 - - 5 8 4
Brassica rapa L. Brassicaceae 15 9 - - - 5 3
Cichorium intybus L. Asteraceae 20 13 - - 6 - -
Colchicum luteum L. Colchicaceae 18 12 - - - - -
Convolvulus arvense L. Convolvulaceae 21 16 - - - - -
Coriandrum sativum L. Apiaceae 12 8 - - 3 5
Cynodia oblonga Rosaceae 11 9 - - - - -
Epilobium hirsutum L. Onagraceae 31 24 - - - - -
Hypericum hookerianum Wight and Arn.
Hypericaceae 23 18 - - - - -
Hypericum perforatum L. Hypericaceae 22 16 - - - - -
Chapter – 3 Determination of pollinator effectiveness among insect visitors
136
Lavatera cashmiriana Cambess. Malvaceae 9 7 - - - - -
Mahonia borealis Takeda Berberidaceae 28 16 - - - - -
Malus domestica Borkh. Rosaceae 17 12 23 - 2 8 - 0
Prunus armeniaca L. Rosaceae 19 15 27 - - - - 0
Punica granatum L. Lythraceae 39 28 - - - - -
Rosa webbiana L. Rosaceae 18 13 23 - - - -
Rosa indica L. Rosaceae 17 8 - - - - -
Sisymbrium irio L. Brassicaceae 18 13 - - - - -
Sternbergia lutea Spreng. Liliaceae 17 13 - - - - -
Chapter – 3 Determination of pollinator effectiveness among insect visitors
137
Table 3.2: Visitation rate of different flower visitors on different plant species
Plant Species
Family
Apis
mellifera
Apis
cerana
Xylocopa
Valga
Bombus
tunicatus
Eristalis
tenax
Andrena
flavipes
Lasioglossum
margenatum
Vanessa
cashmeriansis
Abelia grandiflora Rehd. Caprifoliaceae 4 2 3 2 - - - 1
Actinidia deliciosa Liang and Ferguson.
Actinidiaceae 12 8 - - - - -
Aesculus indica Hook. Sapindaceae 5 6 - - - - -
Alcea rosea L. Malvaceae 6 3 - - - - -
Antirrhinum majus L. Scrophulariaceae
3 - 5 - - - -
Berberis lycium Royle Berberidaceae 3 4 - - 3 - -
Brassica compestris Brassicaceae 3 4 - - 1 2 6
Brassica rapa L. Brassicaceae 4 5 - - - 3 5
Cichorium intybus L. Asteraceae 2 01 - - 1 - -
Colchicum luteum L. Colchicaceae 5 6 - - - - -
Convolvulus arvense L. Convolvulaceae 02 01 - - - - -
Coriandrum sativum L. Apiaceae 04 05 - - 3 5
Cynodia oblonga Rosaceae 03 02 - - - - -
Epilobium hirsutum L. Onagraceae 02 01 - - - - -
Hypericum hookerianum Wight
and
Arn. Hypericaceae 05 02 - - - - -
Chapter – 3 Determination of pollinator effectiveness among insect visitors
138
Hypericum perforatum L. Hypericaceae 04 02 - - - - -
Lavatera cashmiriana Cambess. Malvaceae 05 02 - - - - -
Mahonia borealis Takeda Berberidaceae 03 03 - - - - -
Malus domestica Borkh. Rosaceae 04 03 02 2 2 1
Prunus armeniaca L. Rosaceae 07 08 03 - - - - 2
Punica granatum L. Lythraceae 03 02 - - - - -
Rosa webbiana L. Rosaceae 06 06 02 - - - -
Rosa indica L. Rosaceae 04 03 - - - -
Sisymbrium irio L. Brassicaceae 02 01 - - - - -
Sternbergia lutea Spreng. Liliaceae 04 03 - - - - -
Chapter – 3 Determination of pollinator effectiveness among insect visitors
139
DISCUSSION
The findings of the present study revealed that there is a great deal of
variation in pollen depositing potential of 08 flower visitor species on 25
different plant species and thus reflecting difference in pollinator effectiveness
(Figs. 3.1-3.29; Table 3.1, 3.2). Previous studies have shown that the
pollinating efficacy of flower visitors varies due to difference in pollen
deposition on virgin stigma and their visitation rate (Stephen, 1962; Bohart,
1972). Both are principle factors for checking pollinator validity for a plant
species. Therefore, it becomes necessary to test the insect visitor’s pollen
depositing capacity to differentiate the legitimate pollinators and cheaters for a
particular plant species (Alarcon, 2010; Genini et al., 2010).
A flower of a particular plant species is visited by an array of insect
visitors for feeding, nesting and mating purposes, which in turn helps plant
species in pollen transfer for reproductive success at different rates. The present
study revealed that bigger the size of flower visitor more is the pollen
deposition number on virgin stigma (Figs. 3.1 and 3.29) which coincides with
the studies conducted by Willmer and Finlayson (2014) and Sahli and Connor
(2007). The results revealed that bumbles bees are efficient pollen depositors as
compared to honey bees due to their larger size for Abelia grandiflora (Fig.
3.1); which corroborate with similar results obtained by Ruijter (1997).
The pollination efficacy including pollen depositing ability on virgin
stigma and visitation rate of five different insect visitor species on Malus
domestica revealed that Xylocopa valga deposited large number of pollens but
have less visitation rate as compared to the members of Apis (Fig. 3.17), thus
predicting that Xylocopa valga along with Apis mellifera and A. cerana as
principle pollinators. Similar type of results was obtained by Thomson and
Goodell (2001) on Malus domestica. Among 25 plant species studied, 08 plant
species which include: Actinidia deliciosa, Malus domestica, Brassica
compestris, Brassica rapa, Coriandrum sativum, Cynodia oblonga, Prunus
Chapter – 3 Determination of pollinator effectiveness among insect visitors
140
armeniaca and Punica granatum have horticultural significance and are
economically important crops. Investigating the pollinator efficacy for these
crop species and providing a solution by suggesting alternative pollinator other
than Apis mellifera which is under threat of extinction due to colony collapse
disorder (CCD).
Optimum pollinator selection for a crop plays a vital role (Thomson and
Goodell, 2001). Hence, use of alternative pollinators should be emphasized for
sustainable production in pollinator dependent ecosystems (Stephen, 1962;
Bohart, 1972; Ruijter, 1997; King et al., 2013). The results of pollinator
effectiveness of Apis mellifera and other indigenous insect visitors (Figs. 3.26-
3.29) on different flowering plants are similar to the studies of Paxton (1998).
The present study showed that bees including Xylocopa valga, Bombus
tunicatus, Apis mellifera, A. cerana, Andrena flavipes are the most successful
pollen depositors on virgin stigmas of different flowering plants as compared to
diptera (Eristalis tenax) and butterflies (Vanessa cashmeriansis) (Figs. 3.1 -
3.29). Similar results have been obtained in other parts of the world (Willmer et
al., 1994; Thomson, and Goodell 2001; King et al., 2013). It is found that
larger bees like Xylocopa valga, Apis mellifera and Apis cerana act efficient
pollinators then smaller bees like Lasioglossum margenatum (Sahli and
Connor, 2007; Kandori, 2002). The honey bees are efficient pollen depositors
as well as efficient foragers (Table3.1, 3.2) which is in conformity with
Whelan et al. (2009).
It is pertinent to note that not every flower visitor is the pollinator of a
particular plant species but a pollinator is the flower visitor which deposites the
sufficient number of pollens on the virgin stigma. In other words, all pollinators
are flower visitors but all flower visitors are not pollinators. Without
differentiating the pollinator effectiveness of different flower visitors, it is hard
to think about their conservation and it can misguide our conservational
strategy (Burkle and Alarcon, 2011) and undermine their role of being
Chapter – 3 Determination of pollinator effectiveness among insect visitors
141
sustainers of biodiversity (Ruben et al., 2012). It has been found that larger
flower visitors are efficient pollinators for generalist plant species (Sahli and
Conner, 2007; Kandori, 2002).
Furthermore, the carpenter bee deposited the highest number of pollens
on the virgin stigma but visited flowers less frequently as compared to honey
bees which deposited less pollen but have significant visitation rate. It is not
nesseary that the commom flower visitor can be the best pollinator for a
particular plant species as it is evident from the present study (Table 3.1, 3.2).
For instance, honey bees are the most common foragers but are not efficient as
compared to carpenter bees which are less common foragers on SVD basis
(Fenster, 2004). The behavior of a pollinator at a flower affects the pollinator
effectiveness. Some flower visitors are meant to take nectar only by side
working and are not efficient pollinators where as some are meant to take
pollens by dancing work over anthers and acts as efficient pollinators.
Chapter – 3 Determination of pollinator effectiveness among insect visitors
142
CONCLUSIONS
The present study characterized the pollen depositing ability and
visitation rate of eight (08) different insect visitor species on twenty five (25)
foraging plant species and differentiated efficient pollinators from inefficient
ones. e.g. Xylocopa valga from Eristalis tenax and Vanessa cashmeriansis. The
results reveal that different plant species have specific insect visitors, such as
Xylocopa valga deposits highest number (24) of pollens on the virgin stigma of
Abelia grandiflora followed by Bombus tunicatus (16), Apis mellifera (13),
Apis cerana (07) and visitation rate was found highest for Apis mellifera (04
visits/10 minutes) followed by Xylocopa valga on Abelia grandiflora flowers.
Likewise, Apis mellifera deposited highest (23) number of pollens followed by
Apis cerana (18) and Eristalis tenax (4) on virgin stigmas of Berberis lycium.
The highest visitation rate was recorded for Apis cerana (4) followed by other
three taxa (03 each) on Berberis lycium flowers. Apis mellifera deposited
highest (15) number of pollens followed by Apis cerana (11), Andrena flavipes
(8), Eristalis tenax (5) and Lasioglossum margenatum (4) on virgin stigmas of
Brassica compestris. Lasioglossum margenatum has highest visitation rate (06)
followed by Apis cerana (04), Apis mellifera (03), Andrena flavipes (02) and
(01) of Eristalis tenax on Brassica compestris flowers. Apis mellifera deposited
highest (15) number of pollens followed by Apis cerana (9), Andrena flavipes
(5) and Lasioglossum margenatum (3) on the virgin stigmas of Brassica rapa.
Visitation rate (5 visits/10 minutes) was recorded highest for Lasioglossum
margenatum and Apis cerana followed by Apis mellifera (4) and (03) Andrena
flavipes on Brassica rapa flowers.
Among the Apis genus, A. mellifera had more pollinator effectiveness as
compared to A. cerana both in terms of both pollen deposition as well as
visitation rate on Actinidia deliciosa, Alcea rosea, Cydonia oblonga, Cichorium
intybus, Convolvulus arvense, Epilobium hirsutum, Hypericum hookerianum,
H. perforatum, Lavatera cashmiriana, Mahonia borealis, Punica granatum,
Chapter – 3 Determination of pollinator effectiveness among insect visitors
143
Rosa indica, Sisymbrium irio, Sternbergia vernalis. It was found that Apis
mellifera had high pollen deposition ability than A. cerana and A. cerana had
highest visitation rate than A. mellifera on Aesculus indica, Berberis lyceum,
Brassica compestris, B. rapa, Colchicum luteum, Coriandrum sativum, Malus
domestica, Prunus armeniaca, Rosa webbiana. A comparative analysis of
pollen deposition by Apis mellifera and Apis cerana on different flowering
plants revealed that Apis mellifera deposited highest number of pollens than A.
cerana on all plant species. Xylocopa valga deposited the highest number of
pollens on virgin stigmas followed by Apis mellifera and A. cerana. Thus
Xylocopa valga serves as best pollen depositing insect visitor followed by
Bombus tunicatus, Apis mellifera, A. cerana, Andrena flavipes, Lasioglossum
marginbatum and finally Eristalis tenax is revealed from above results.
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Chapter – 4 Characterization of generalized and specialized plant insect interactions
151
INTRODUCTION
Flower insect interaction web is the life pool of two different life forms
which interact with each other in order to continue their race and helps to
produce ecosystem outcome. The diversified lifeform (i.e. angiosperms) is
because of the interaction web with another diversified lifeform (i.e.
pollinators). In this interactive web, different workers have different interacting
magnitude, thus resulting into generalized and specialized plant pollinator
interactions. Among the plant pollinator interacting web, specialized plant
pollinator interaction is vulnerable to pollinator deficits, as most of plant
pollinator web is generalized as compared to specialized because most of
flower visitors have wider range of flower choice (Petanidou et al., 2006).
Even though certain plant pollinator interactions are specialized due to
selective trait morphology of flowers which restrict access to only particular
pollinators (Fenster et al., 2004) and eliminating other flower visitors which are
meant to rob the floral reward (Alarcon, 2010). In addition to significance of
interaction web to these two interacting forms, it serves as an essential
ecosystem service and a base to enhance the quality and quantity of ecosystem
outcome.
In an ecosystem, a flowering plant species interacts with different flower
visitor species and vice-versa. It has, therefore become imperative to study the
nature of plant pollinator interaction of an ecosystem (Corbet, 2000). Recent
studies have shown that majority of plant pollinator interactions are generalized
i.e. those plant species which are visited by a diversified pollinator fauna for
their food as pollen and nectar (Kandori, 2002; Waser et al., 1996). This has
given rise to the debate about relative frequency of generalization of plant
pollinator interaction (Waser et al., 1996) and also how generalization should
be characterized (Herrera, 2005) and can be helpful in understanding the plant
pollinator web (Memmott and Waser, 2002). These two types of interactions
are biologically two sides of the same coin which is known as generalization-
Chapter – 4 Characterization of generalized and specialized plant insect interactions
152
specialization continuum (Johnson and Steiner, 2000). Plant pollinator
interaction is the most valuable service of ecosystem, whose value is
incalculable (Kearns et al., 1998). Categorization of flower insect interactions
into the generalization and specialization may guide towards their conservation
and sustainable use.
Plant pollinator interaction has been categorized into generalization and
specialization on the basis of number of different flower visitor taxa visiting a
particular foraging plant species (Sahli and Conner, 2006). In understanding
generalized interaction, scientists have used both quantitative (visitation rates)
and qualitative attributes (pollinator effectiveness) of different flower visitors
on a particular plant species (Johnson and Steiner, 2000). Pollinator
effectiveness and pollinator visitation frequency cumulatively gives the
pollinator importance, a measure of pollinator contribution for a plant species
(Inouye et al., 1994). There is great dependence of reproductive success of
flowering plants and crop plants on pollinators which caused concern that
pollinator decline will cause decline in ecosystem outcome (Kearns et al.,
1998).
REVIEW OF LITERATURE
Sahli and Conner (2006) studied the plant pollinator interaction for 17
plant species by studying visitation richness and found that visitation richness
for all ranged from 2-45 pollinator genera and two third of plant species
showed more specialized interaction having visitation richness less than 07 taxa
were classified as specialized plants and others have greater than 07 visitation
rich were classified as generalized plants. Sahli and Conner (2007) studied the
visitation, effectiveness and efficiency of 15 genera of flower visitors on wild
radish (Raphanus raphanistrum) and found that the plant serves as the member
of generalized pollination system. Fang et al. 2012 studied the pollination in
Chapter – 4 Characterization of generalized and specialized plant insect interactions
153
winter flowering fruit (Eriobotrya japonica) and found that insect visitation
was higher in early winter and in late winter insect visitation ceased but two
species of passerine bird pollinate the tree in extreme cold condition reflecting
generalized type of pollination system in Eriobotrya japonica due to wide
range of visited pollinator taxa.
Gomez and Zamora (1999) studied the pollination of a crucifer shrub
Hormathophylla spinosa in Spain and found that a large number of pollinator
taxa including pollenivorous as well as nectarivorous were visiting the above
mentioned plant species for pollen or nectar which reflects the generalized type
of pollination system in Hormathophylla spinosa. Kyra and Scott (2014)
studied pollination systems in 26 taxa of Oenethera and found that both types
of pollination systems generalized as well as specialized systems were present
in above mentioned taxa, but more systems were found as specialized one.
Muchhala et al. (2008) studied pollination system in Aphelandra acanthus in
the tropics and found that birds, bats were the effective pollinator fauna visiting
above mentioned plant reflecting generalized pollination system. Kwapong et
al. (2013) studied the pollination of cowpea (Vigna unguiculata) and found that
a large number of flower visitor species were foraging the particular plant for
pollen and nectar reflecting the generalized system of pollination.
Vazquez et al. (2004) found that pollinator communities comprises of
different groups highly specialized pollinators which are rare, many moderate
specialized pollinators and generalist pollinators. Bascompte and Jordano
(2007) studied the ecology of pollinators, flowers and found that flower
pollinator webs are asymmetrical due to interaction of specialized pollinators
with generalized flowers. Paray et al. (2014) studied the interaction of apple
with flower visitors and found that 15 species of flower visitors belonging to 8
genera and 4 families were interacting with the Malus domestica. Ganie et al.
(2013) studied the diversity of flower visitors of Malus domestica and found
that 16 species of flower visitors belonging to 11 families and 3 orders
Chapter – 4 Characterization of generalized and specialized plant insect interactions
154
Hymenoptera, Diptera and Lepidoptera were foraging the apple flowers for
their food in the Kashmir Valley.
A recent upsurge of generalization-specialization continuum
measurement shown by plant pollinator interactions is found (Johnson and
Steiner, 2000) due to increasing recognition of specialization in plant pollinator
web (Pellmyr, 2002). Petanidou and Lambron (2005) reviewed the plant
pollinator interaction by plant pollinator web approach and found that different
authors have studied the ecological aspects of plant pollinator interaction
including generalization and specialization interactions among them. It
becomes imperative for pollination biologist to study the relationship between
pollinators and their foraging plant species and nature of pollination web
(Corbet, 2000). Jiliberto et al., (2009) characterises the structure of plant
pollinator web in the temperate rain forest south Chile and found that trees and
hymenopteran pollinators are pivotal for maintains of plant pollinator web
structure of Chile and reported that tree species should be focal plants for
conservational purposes as Burkle (2013) found that decrease in foraging plant
species may lead to extinction of specialized pollinator taxa. Memmott (1999)
reviewed that plant pollinator interaction literature is dominated by the
presence of specialization in the interaction and showed that majority of plant
pollinator interactions are embedded in the plant pollinator web.
MATERIAL AND METHODS
By using random sampling design, pollinators were observed on studied
plants from dawn to dusk throughout the study period during 2014-2015.
Measurement of generalization and specialization in plant pollinator interaction
is the perception of generalization and specialization of plant species and not of
pollinator species. Plant generalization and specialization is defined by various
authors (Armbruster et al., 2000; Johnson and Steiner, 2000; Gomez, 2002;
Kay and Schemske, 2004; Jiliberto, 2009; Kyra and Scott, 2014) but most
Chapter – 4 Characterization of generalized and specialized plant insect interactions
155
accurately was defined by Ollerton and Cranmer 2002 that plant pollinator
generalization and specialization are defined on the basis of number of
pollinator taxa (genera) interacting with the particular foraging plant species
and was quantified by visitor-centred approach (Herrera 1988; Mahy et al.,
1998; Dicks et al., 2002). Interactions where a plant species is foraged by more
than one insect visitor genera were classified as generalized plant pollinator
interactions where as interactions in which a foraging plant species was foraged
by only one pollinator genera were classified as specialized plant pollinator
interactions. Pollinator generalization and specialization was also measured by
its range of foraging plant species. Pollinator foraging a large number of plant
species was classified as generalist and pollinator foraging a particular plant
species was classified as specialist.
RESULTS
The analysis of present study revealed that out of 215 plant pollinator
interactions 134 (62%) are generalized whereas 81 (38%) are specialized (Fig.
4.1; Table 4.1 and photoplates 4.1-4.5). These two sets of interactions involve
215 foraging plant species and 70 different flower visitor species (Table 4.1).
Figure 4.1: Percentage of generalized and specialized plant pollinator
interactions.
Generalized
134,62%
Specialized
81,38%
Chapter – 4 Characterization of generalized and specialized plant insect interactions
156
Among 215 different foraging plant species, 03 plant species are foraged
by 01 flower visitor species, 79 plant species foraged by 2 species of flower
visitors, 22 plant species by 3 flower visitor species, 20 plant species by 4
flower visitor species, 11 plant species by 5 flower visitor species, 11 plant
species by 6 flower visitor species, 9 plant species by 7 flower visitor species,
12 plant species by 8 flower visitor species, 12 plant species by 9 flower visitor
species, 9 plant species by 10 flower visitor species, 5 plant species by 11
flower visitor species, 2 plant species by 12 flower visitor species, 3 plant
species by 13 flower visitor species, 6 plant species by 14 flower visitor
species, 1 plant species by 15 flower visitor species, 3 plant species by 16
flower visitor species, 3 plant species by 21 flower visitor species, 2 plant
species by 22 flower visitor species and 2 plant species by 23 flower visitor
species (Fig. 4.2).
Figure 4.2: Number of flower visitor species interacting with number of plant
species.
3
79
22 20
11 11 912 12 95236
13 3 2 2
12345678910 11 12 13 14 15 16
21 22 23
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
Number of plant species Number of flower visitor species
Chapter – 4 Characterization of generalized and specialized plant insect interactions
157
Plant generalization specialization continuum
Plant generalization specialization continuum revealed that Oxytropis
cashmeriana is foraged by Macroglossum nycteris only, Datura stramonium
and Bidens tripartita by Episyrphus balteatus only. Actinidia deliciosa, Albizia
julibrissin, Arctium lappa, Carya illinoinensis, Chenopodium album, Cucurbita
maxima, Cucumis sativus are foraged by two species of pollinators. Aesculus
indica, Ailanthus altissima, Cichorium intybus, Convolvulus arvense are
foraged by 3 species of pollinators. Prunus avium, Prunus persica and
Capsella bursa pastoris are foraged by 16 flower visitor species. Trifolium
pratense and T. repense are foraged by 21 species belonging to 10 genera of
flower visitors. Taraxacum officinale are foraged by 21 species belonging to 12
genera of insect visitors. Pyrus communis and Pyrus pyrifolia are foraged by
22 species belonging to 11 genera of flower visitors and Malus domestica and
Prunus domestica are foraged by the maximum number (23) of flower visitor
species belonging to 12 genera. The scale on generalization specialization
continuum of others plant species are shown in Fig. 4.3 and in Table 4.1. Out
of 79 foraging plant species which are visited by two species of flower visitors
75 plant species are foraged only by Apis cerana and A. mellifera. Pinus
wallichiana was foraged by Bombus tunicatus and Bombus trifasciatus, Aster
thomsonii was foraged by Lassioglossum margenatum, Lassioglossum
himalayense, and Galinsoga parviflora by Episyrphus balteatus, Frankliniella
sp. (Table 4.1).
Chapter – 4 Characterization of generalized and specialized plant insect interactions
158
Specialization (81 plant species) Generalization (134 plant species)
(0 1) D a t u r a stram o niu m
Bid en s trip a rtita (0 1) (0 1) O xyt ro p i s c a s h m eri a n a
Actin idia d e l icio s a (0 2) ( 0 2) A lb izia ju libri s s in
Arctiu m lappa (02 ) (0 2 ) C a ry a illin o in e n s i s
Che n o po d ium albu m (02 ) (0 2 ) C u c u r b ita m ax im a
Cucum is sa tivus (0 2 ) (0 2 ) Lo n icer a japo n ica
Ae s c u l u s in d ica (0 3 ) (0 3) A i la nth u s a l tis s im a
Ci c h o rium intyb us (03 ) (0 3 ) C o n vo lvulus a rv e n se
Berbe r is lyc i u m (0 4 ) (0 4 ) Co lchicu m lute um
Eschscho lzia c a lifo rn ica (04 ) (0 5 ) C a stane a sa tiv a
Fr a g ar ia an a n a ssa (0 5
)
(0 5) G l a d io lus h ortu la n u s
M en tha l o n g ifo lia (0 5 ) (0 6 ) Calend u la officin alis
Coreo p s is la n c e ola ta ( 0 6 ) (0 6 ) Foe n icu lu m vu lgare
Ple ctran thus rugo s u s (0 6 ) (0 7 ) B u d d leja d a vidii
Conium m a c ula tum (0 7 ) (0 7 ) E rio botyri a japo n ica
Iris de c or a (0 7 ) (0 8) A b el ia g r a n d ifl o r a
Age r a tu m ho u s tonia nu m (0 8) (0 8) A lliu m c e pa
Anthe m is c o tula (08) (08 ) A n t irrh i n u m m a jus
Ca mp sis g r a n d iflo r a (0 8)
(0 9) C a r d u u s e d elb e rg ii
Po te n tilla rep ta ns ( 0 9) (0 9) B e r g e nia lig u lata
Astraga lu s ca ndo lle an u s (10 ) ( 1 0 ) Astr a g a lus g r a h am ianu s
Be llis pe re n n is (1 0) (10 ) Cir s i u m falc o n e ri
Cir s ium arv en se (1 1 ) (1 1 ) Cirsiu m v u l g a r e
Ranu n c u lu s la e tus (1 1) (11 ) Tagetes pa tula
Tu lip a ste llata ( 1 1 ) (1 2) B ra s s ica c o mp e s tris
Ch rys a n the m u m co ro n a rium (1 2 ) (1 3 ) Veron ica arve n s is
Ve r o n ica persic a (1 3 )
(1 4) B r a s s ica r a p a
Ce rcis ca n a d ens is (1 4) (1 3) C o rian d ru m sa tivu m
Li n d e lofia lon g ifl o ra (1 4 ) (14 ) Rosa c an ina
Ro s a bru non ii ( 1 4) (1 5) Z inn ia e leg an s
Pru nus aviu m (1 6) (1 6) P r u n u s pe r s ica
Tar a x a c u m of fic i n a le (2 1 ) (2 1 ) Tr ifo lium prate ns e
Trifoliu m rep e n s (21 ) ( 2 2 ) P yru s com mu nis
Pyru s pyrifo lia (2 2
)
(2 3) M alus d o m es tic a
Prunu s d o m es tic a (2 3)
Figure 4.3 Generalization specialization continuum of Foraging plant species
Numbering (1-23) = Number of pollinator taxa
Pollinator generalization specialization continuum
Pollinator generalization specialization continuum revealed that among
70 different flower visitor species Apis cerana is the most generalist pollinator
visitor foraging 205 plant species followed by Apis mellifera foraging 204 plant
species Eristalis tenax (41), Bombus trifasciatus (38), Bombus tunicatus (37),
Lasioglossum margenatum (35), Lassioglossum himalayense (30),
Lassioglossum nursei (29), and Episyrphus balteatus, Lassioglossum
rugolatum, Lassioglossum polyctor, Lasioglossum sublaterale, Lasioglossum
leucozonium by 26, Pieris brassicae forages 25 plant species, Xylocopa valga,
Xylocopa violacea forages 24 plant species, Formica sp., Andrena flavipes
Chapter – 4 Characterization of generalized and specialized plant insect interactions
159
forages 22 plant species, Vanessa cashmiriensis forages 21 plant species,
Bombus simillmus, Eoseristalis cerealis forages 19 plant species, Vanessa
cardui forages 14 plant species, Scathophaga stercoraria forages 13 plant
species, Bombus rufofasciatus, Bombus pyrosoma forages 12 plant species,
Bombus asiaticus, Eristalinus taeniops and Megachile rotundata forages 10
plant species, Bombus miniatus forages 9 plant species, Andrena patella,
Andrena floridula forages 8 plant species, Ceratina hieroglyphica, Hesperia
comma forages 7 plant species, Frankliniella sp., Sphaerophoria bengalensis,
Bibio marci, Plecia nearctica, Cetonia aurata forages 6 plant species,
Eristalinus aeneu, Metasyrphus bucculatus, Musca domestica, Polyommatus
icarus, Danais chrysippus forages 5 plant species, Sphaerophoria macrogaster,
Issoria lathonia forages 4 plant species, Mordella sp., Amegilla fallax, Vespa
velutina, Pyrrhocorus sp., Colias philodice, Pontia daplidice forages 3 plant
species, Ceratina propinqua, Sphecodes lasimensis, Sphecodes tantalus,
Coccinella magnifica, Mylabris pustulata, Scolia nobilitata, Halictus
constrictus, Colias fieldi, Macroglossum nycteris forages 2 plant species,
Macroglossum nycteris, Heriades sp., Scolia hauseri, Scolia soror, Scoliinae
proscoliinae, Spilomyia sulphurea forages only one plant species. (Figs. 4.4-
4.8 and Table 4.1).
Chapter – 4 Characterization of generalized and specialized plant insect interactions
160
Specialization Generalization
Figure 4.4 Generalization specialization continuum of pollinator species
(01 ) Spilom yia sulph ur ea
Scoliina e proscoliin ae (01) (01) Scolia sor or
Scolia hauseri (01) (01) Heriades sp
.
Eristalod es paria (01) (02) M acroglossum ny cteris
Colias fieldi (02) (02) Ha lictu s constrictus
Scolia nobilitata (02) (02) M y labris pustulata
Coccinella m agnific a (0 2) (02 ) S phecod es ta ntalus
Sphecodes la simensis (02) (02) Ceratina propinqu a
Pontia daplidice (03) (0 3) Colia s philo dice
Pyrrhocorus sp. (0 3) (03) Vespa velutina
Am egilla fallax (03) (03) Mordella sp.
Issoria lathonia (04) (04) Sphaerophoria m ac ro ga ster
Danais ch ry sippus (05) (05) Polyo m matus ic ar us
Musca domestica (05 ) (05) Metasyrphus buccu latus
Eristalinu s aeneu (0 5) (06) Ceton ia aurata
Plecia nearctica (06) (06) Bibio marci
Sphaero phoria bengalensis (06) (06 ) Fr an kliniella sp.
Hesperia c om ma (0 7) (07) C eratina hiero glyphica
Andrena flor idula (08) (0 8) Andren a patella
Bomb us m iniatus (09) (1 0) Er ista linus tae niops
Bomb us a siaticus (1 0) (12 ) B om bus py rosoma
Bombus rufofascia tus (12) (1 3) Scat ho ph ag a stercora ria
Vanessa ca rdui (14) (19) Eo se ristalis cere alis
Bomb us simillm us (19) (21 ) Vanessa ca sh miriensis
Andrena flavipes (2 2) (22 ) Formica sp.
Xylocop a violacea (2 2) (22 ) X ylocopa valg a
Pieris brassic ae (25) (26) Las ioglo ssum leucozonium
Lasioglo ssum subla ter ale (26) (26) Lassioglossum po lycto r
Las sioglossum rugolatum (26) (26) Episyrphus balte atus
Lassioglossum nursei (29) (30) Lassioglossum himalayense
Lasioglo ssum margenatum (35) (37) Bombus tunic atus
Bomb us trifasciatus (38) (41) Er ista lis tenax
Me ga chile rotundata (1 09) (204 ) A pis m ellifer a
Apis cerana (205)
Numbering = Number of foraging plant species
Figure 4.5: Generalization in pollinator species
0
50
100
150
200
250
Number of foraging
plant species
Pollinator species
Chapter – 4 Characterization of generalized and specialized plant insect interactions
161
Figure 4.6: Generalization in pollinator species
Figure 4.7: Generalization in pollinator species
0
2
4
6
8
10
12
14
16
18
20
Number of foraging plant
species
Pollinator species
0
1
2
3
4
5
6
Pollinator species
No. of foraging plant
species
Chapter – 4 Characterization of generalized and specialized plant insect interactions
162
Figure 4.8: Generalization and specialization of pollinator species.
Plant pollinator web and its interacting pillars
The results reveal that the plant pollinator web of Kashmir Himalaya is
constructed by four different interacting life forms viz. generalized plant
species, specialized plant species, generalized pollinator species and
specialized pollinator species.
Generalized plant species
Generalized plant species are those flowering plant species which are
visited by more than one genus of flower visitors. Among 215 flowering plant
species 134 plant species acts as generalists viz. Malus domestica, Prunus
domestica, Pyrus communis, Pyrus pyrifolia, Taraxacum officinale, Trifolium
pratense, Trifolium repens, Prunus avium, Prunus persica, Brassica rapa etc.
are foraged by more than one genus of flower visitors (Table 4.1)
0
0.5
1
1.5
2
2.5
Number of foraging plant
species
Pollinator species
Chapter – 4 Characterization of generalized and specialized plant insect interactions
163
Specialized plant species
Specialized plant species are those flowering plant species which are
visited by only one genus of flower visitors. 81 plant species viz. Actinidia
deliciosa, Albizia julibrissin, Arabis glabra, Arctium lappa, Aster thomsonii,
Bidens tripartita, Carya illinoinensis, Centaurea cyanus, Chenopodium album,
Crepis tectorum, Cucumis melo, Cucumis sativus etc are visited by one or two
species of same genus (Table 4.1).
Generalized pollinator species
Generalized pollinators are those which forage the maximum number of
flowering plants for their food as pollen and nectar viz. Apis cerana forages
205 flowering plants followed by Apis mellifera (204), Megachile rotundata
(109), Eristalis tenax (41), Bombus trifasciatus (38), Bombus tunicatus (37),
Lasioglossum margenatum (35), Lassioglossum himalayense (30),
Lassioglossum nursei (29), and Episyrphus balteatus, Lassioglossum
rugolatum, Lassioglossum polyctor, Lasioglossum sublaterale, Lasioglossum
leucozonium by 26 etc. (Figs. 4.4-4.7 and Table 4.1).
Specialized pollinator species
Specialized pollinators are those which forage only one flowering plant
for their food either pollen or nectar viz. Eristalodes paria and Spilomyia
sulphurea forages on Ageratum houstonianum, Heriades sp. forages on
Chrysanthemum coronarium, Scolia hauseri, Scolia soror, Scoliinae
proscoliinae forages on Mentha arvensis (Fig. 4.7 and Table 4.1) shaping
three types of specialized plant pollinator interaction.
These four interacting pillars shapes plant pollinator interaction in four
different ways. Present study revealed that 130 interactions are found to exist
between generalized plant species and generalized flower visitor species viz.
interaction between Abelia grandiflora and Apis cerana or Apis mellifera. 82
interactions are are found to exist between specialized plant and generalized
Chapter – 4 Characterization of generalized and specialized plant insect interactions
164
flower visitor viz. interaction between Actinidia deliciosa and Apis cerana or
Apis mellifera. 6 interactions are found to exist between generalized plant
species and specialized flower visitor viz. interaction between Ageratum
houstonianum and Spilomyia sulphurea and non interaction was found existing
between specialized plant species and specialized flower visitor species in the
present study (Fig. 4.9 and Table 4.1).
Flowering period of foraging plants
Flowering calendar of studied flora ranges from one to eight months,
longest flowering period was recorded for Trifolium pratense and Trifolium
repense (08 months) followed by Rosa indica, Taraxacum officinale, Veronica
arvensis, Veronica arvensis and Solanum nigrum 7 months and shortest
flowering period was recorded one month for Castanea sativa, Carya
illinoinensis, Cynodon dactylon, Forsythia viridissima etc.(table 4). It was
found that Trifolium pratense and Trifolium repense having the longest
flowering period (08 months) harbour maximum number of flower visitor
species followed by Taraxacum officinale having flowering period for 7
Plant pollinator interaction
Generalized
Plant
generalized
Pollinator
Interaction
(130)
Generalized
plant
specialized
pollinator
interaction
(6)
Specialized
plant
Generalized
pollinator
interaction
(82)
Specialized
plant
specialized
pollinator
interaction
(0)
Figure 4.9 Types of interactions
Chapter – 4 Characterization of generalized and specialized plant insect interactions
165
months harbour 21 species of flower visitors, Veronica arvensis and Veronica
persica (07 months) harbour 13 species of flower visitors (Table 4.1)
Among 215 foraging plant species, 121 (56%) are native where as 94
(44%) are exotic to Kashmir Himalaya (Fig. 4.10; Table 4.1)
Figure 4.10: Percentage contribution of foraging plant species in terms of
native and exotic species.
Native
121,56%
Exotic
94,44%
Chapter – 4 Characterization of generalized and specialized plant insect interactions
166
Table 4.1: Inventory of plant pollinator generalization and specialization
Plant species Family Origin
Flowering
period
Number of
months Flower visitors
Number of
taxa
Type of
interaction
Abelia grandiflora
Rehd. Caprifolaceae Exotic Aug-Oct 3
Xylocopa valga, Xylocopa violacea, Bombus tunicatus,
Bombus trifasciatus, Apis cerana, Apis mellifera,
Eoseristalis cerealis, Hesperia comma.
5 G
Abelmoschus
esculentus Moench Malvaceae Exotic
Xylocopa valga, Xylocopa violacea, Bombus tunicatus,
Bombus trifasciatus, Apis cerana, Apis mellifera. 3 G
Aconitum laeve
Royle Ranunculaceae Native
Apis cerana, Bombus miniatus, Bombus pyrosoma,
Bombus asiaticus, Bombus rufofasciatus,Bombus
trifasciatus, Bombus simillmus, Bombus tunicatus
2 G
Actinidia deliciosa
Liang and Ferguson.
Actinidiaceae Exotic May, June 2 Apis cerana, Apis mellifera 1 S
Aesculus indica
Hook. Sapindaceae Native
May 1 Apis cerana, Apis mellifera, Episyrphus balteatus, 2 G
Ageratum
houstonianum Mill. Asteraceae Exotic July-Oct 4
Apis cerana, Apis mellifera, Eristalis tenax, Eoseristalis
cerealis, Sphaerophoria bengalensis, Athalia proxima,
Spilomyia sulphurea, Eristalodes paria.
7 G
Agrimonia pilosa
Ledeb. Rosaceae Native
Apis cerana, Apis mellifera. Lasioglossum margenatum. 2 G
Ailanthus altissima
Swingle Scrophulariaceae
Exotic May, June 2 Apis cerana, Apis mellifera, Eristalis tenax 2 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
167
Albizia julibrissin
Durazz. Fabaceae Exotic June-July 2 Apis cerana, Apis mellifera. 1 S
Alcea rosea L. Malvaceae Exotic June-Aug 3
Apis cerana, Apis mellifera, Bombus trifasciatus, Bombus
tunicatus. 2 G
Allium cepa L.
Alliaceae Exotic May, June 2
Apis cerana, Apis mellifera, Megachile rotundata,
Ceratina hieroglyphica, Lassioglossum margenatum,
Eristalis tenax,Eristalinus aeneu, Mordella sp.
7 G
Allium sativum L.
Alliaceae Exotic May, June 2
Apis cerana ,Apis mellifera,
Megachile rotundata,
Ceratina hieroglyphica, Lassioglossum margenatum,
Eristalis tenax,Eristalinus aeneu, Mordella sp.
7 G
Anemone obtusiloba
Don Ranunculaceae Native
May, June 2
Apis cerana, Apis mellifera, Lasioglossum nursei,
Lasioglossum margenatum. 2 G
Anemone
tetrasepala Royle Ranunculaceae Native
May, June 2
Apis cerana, Apis mellifera, Lassioglossum nursei,
Lasioglossum margenatum. 2 G
Anthemis cotula L.
Asteraceae Exotic May-Sept 5
Apis cerana, Apis mellifera, Polyommatus icarus,
Eristalis tenax, Eristalinus taeniops, Episyrphus
balteatus, Frankliniella sp., Scathophaga stercoraria.
7 G
Antirrhinum majus
L. Scrophulariaceae
Exotic June-Oct 5
Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Bombus tunicatus,
Bombus trifasciatus,
Ceratina hieroglyphica, Ceratina propinqua.
4 G
Arabis glabra
Bernh. Brassicaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Arctium lappa L. Asteraceae Exotic May-July 3 Apis cerana, Apis mellifera 1 S
Aster thomsonii Asteraceae Native
June-Aug 3 Lassioglossum margenatum, Lassioglossum himalayense,
1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
168
Clarke
Astragalus
candolleanus Fabaceae Native
March-May
3
Apis mellifera, Apis cerana, Amegilla fallax, Formica,
Andrena flavipes, Andrena patella, Andrena floridula,
Xylocopa valga, Xylocopa violacea, Metasyrphus
bucculatus.
6 G
Astragalus
grahamianus Benth.
Fabaceae Native
March-May
3
Apis mellifera, Apis cerana, Amegilla fallax, Formica,
Andrena flavipes, Andrena patella, Andrena floridula,
Xylocopa valga, Xylocopa violacea, Metasyrphus
bucculatus.
6 G
Bellis perennis L.
Asteraceae Exotic Feb- May 4
Apis cerana, Apis mellifera,
Andrena flavipes,
Lassioglossum margenatum, Eoseristalis cerealis,
Sphaerophoria bengalensis, Sphaerophoria macrogaster,
Scathophaga stercoraria, Musca domestica.
7 G
Berberis lycium
Royle Berberidaceae Native
April, May
2 Apis cerana, Apis mellifera,
Eristalinus taeniops, Eristalis
tenax, 3 G
Bergenia ligulata
Engl. Saxifragaceae Native
May- July 3
Apis cerana,
Apis mellifera, Bombus miniatus, Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus,
Bombus trifasciatus, Bombus simillmus,Bombus tunicatus
2 G
Bidens tripartita L. Asteraceae Exotic
Episyrphus balteatus 1 S
Brassica compestris
Brassicaceae Exotic April, May
2
Apis cerana,
Apis mellifera, Lassioglossum
nursei,Lassioglossum himalayense,
Lassioglossum
rugolatum,
Lassioglossum polyctor, Lasioglossum
margenatum, Lasioglossum sublaterale,
Lasioglossum
leucozonium,Andrena flavipes,
Episyrphus balteatus,
5 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
169
Vanessa cardui,
Brassica rapa L.
Brassicaceae Exotic April, May
2
Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale, Lasioglossum leucozonium, Andrena flavipes,
Episyrphus balteatus, Vanessa cardui,
6 G
Buddleja davidii
Franch. Scrophulariaceae
Exotic July-Aug 2
Apis cerana, Apis mellifera, Vanessa cashmiriensis,
Vanessa cardui, Pieris brassicae, Eristalis tenax,
Eoseristalis cerealis.
5 G
Calendula officinalis
L. Asteraceae Exotic July-Aug 2 Apis cerana, Apis mellifera, Frankliniella sp
., Bombus
tunicatus, Bombus trifasciatus, Bombus simillmus, 3 G
Caltha alba
Cambess. Ranunculaceae Native
May-Sept 5
Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium,
2 G
Campanula
cashmeriana Royle
Campanulaceae Native
Apis cerana, Apis mellifera, Bombus trifasciatus, Bombus
rufofasciatus, Bombus tunicatus, Bombus asiaticus,
Bombus pyrosoma.
2 G
Campanula
rotundifolia L. Campanulaceae Native
Apis cerana, Apis mellifera, Bombus trifasciatus, Bombus
rufofasciatus,
Bombus tunicatus, Bombus asiaticus,
Bombus pyrosoma.
2 G
Campsis grandiflora
Schum. Bignonaceae Exotic July-Sept 3 Xylocopa valga, Xylocopa violacea, Bombus tunicatus,
Bombus trifasciatus, Apis cerana,
Apis mellifera, Vespa
5 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
170
velutina, Formica
Capsella bursa-
pastoris
Brassicaceae Native
March-June
4
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor
,
Lasioglossum margenatum, Lasioglossum sublaterale
,
Lasioglossum leucozonium, Episyrphus balteatus,
Megachile rotundata, Sphaerophoria fatarum,
Sphaerophoria bengalensis,
Sphaerophoria macrogaster,
Eristalinus aeneu, Andrena flavipes, Andrena patella,
Scathophaga stercoraria.
7 G
Carduus edelbergii
Rech.f. Asteraceae Native
May, June 2
Apis cerana, Apis
mellifera, Ceratina hieroglyphica,
Eristalis tenax, Vanessa cashmiriensis, Xylocopa valga,
Xylocopa violacea, Andrena flavipes, Pieris brassicae,
7 G
Carya illinoinensis
Koch Juglandaceae Exotic May 1 Apis cerana, Apis mellifera 1 S
Castanea sativa
Mill. Fagaceae Exotic June 1
Apis cerana, Apis mellifera, Eristalis tenax, Eoseristalis
cerealis. Eristalinus taeniops, 4 G
Centaurea cyanus L.
Asteraceae Exotic May, June 2 Apis cerana, Apis mellifera 1 S
Centaurea iberica
Spreng. Asteraceae Native
Apis cerana, Apis mellifera, Formica, Andrena patella,
Megachile rotundata, 4 G
Cercis canadensis L.
Fabaceae Exotic April, May
2
Apis cerana ,Apis mellifera, Xylocopa valga ,
Xylocopa
violacea, Vanessa cashmiriensis, Vanessa cardui,
Cetonia
aurata, Lassioglossum nursei
, Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
5 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
171
sublaterale, Lasioglossum leucozonium
Chenopodium album
L. Amaranthaceae Exotic June-Sept 4 Apis cerana, Apis mellifera 1 S
Chrysanthemum
coronarium Spach
Asteraceae Exotic May 1
Apis cerana,
Apis mellifera, Sphecodes lasimensis,
Sphecodes tantalus, Coccinella magnifica, Andrena
flavipes, Lassioglossum nursei, Eoseristalis cerealis,
Eristalis tenax, Eristalinus aeneu, Eristalinus taeniops,
Heriades,
9 G
Cichorium intybus
L. Asteraceae Native
June-Oct 5 Apis cerana, Apis mellifera, Episyrphus balteatus. 2 G
Cirsium arvense
Scop. Asteraceae Native
May-Sept 5
Apis cerana, Apis mellifera, Eristalis tenax, Eristalinus
taeniops, Bombus tunicatus, Bombus trifasciatus, Danais
chrysippus, Scolia
hirta. Andrena flavipes, Xylocopa
valga, Xylocopa violacea.
8 G
Cirsium falconeri
Petr. Asteraceae Native
May-Sept 5
Apis cerana, Apis mellifera, Hesperia comma, Cetonia
aurata, Bombus tunicatus, Bombus trifasciatus, Danais
chrysippus, Scolia sp.,
Xylocopa valga, Xylocopa
violacea.
7 G
Cirsium vulgare
Ten. Asteraceae Native
May-Sept 5
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus, Pieris brassicae, Ceratina hieroglyphica,
Ceratina propinqua, Danais chrysippus, Scolia sp
.,
Xylocopa valga, Xylocopa violacea.
7 G
Clematis montana
DC. Ranunculaceae Native
Apis cerana, Apis mellifera, Cetonia aurata,
Bombus
miniatus, Bombus pyrosoma,
Bombus asiaticus, Bombus
3 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
172
rufofasciatus, Bombus trifasciatus, Bombus simillmus
,
Bombus tunicatus,
Colchicum luteum L.
Colchicaceae Native
Feb., March
2
Apis cerana, Apis mellifera, Halictus propinquus,
Scathophaga stercoraria. 3 G
Conium maculatum
L. Apiaceae Exotic May, June 2
Apis cerana,
Apis mellifera, Eoseristalis cerealis,
Eristalis tenax, Coccinella magnifica, Pyrrhocorus sp
.,
Athalia proxima,
6 G
Convolvulus arvense
L. Convolvulaceae Native
May-Sept 5 Apis cerana, Apis mellifera, Episyrphus balteatus 2 G
Coriandrum sativum
L.
Apiaceae Exotic May-July 3
Apis cerana,
Apis mellifera, Andrena flavipes, Eristalis
tenax, Tacnidae, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale, Lasi
oglossum leucozonium, Scathophaga
stercoraria.
6 G
Coreopsis
lanceolata L. Exotic
Apis mellifera, Apis cerana, Lasioglossum margenatum,
Megachile rotundata, Andrena flavipes, Eristalis tenax. 5 G
Crepis tectorum L. Asteraceae Native
May, June 2 Apis cerana, Apis mellifera 1 S
Cucumis melo L. Cucurbitaceae Exotic July-Sept 4 Apis cerana, Apis mellifera 1 S
Cucumis sativus L. Cucurbitaceae Exotic July-Sept 4 Apis cerana, Apis mellifera 1 S
Cucurbita maxima
Duchesne Cucurbitaceae Exotic July-Nov 5 Apis cerana, Apis mellifera 1 S
Cucurbita pepo L. Cucurbitaceae Exotic July-Nov 5 Apis cerana, Apis mellifera 1 S
Cynodia oblonga Rosaceae Exotic April, May
2 Apis cerana, Apis mellifera 1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
173
Cynodon dactylon
Pers. Poaceae Native
May 1 Apis cerana, Apis mellifera 1 S
Cynoglossum
glochidiatus Wall Boraginaceae Native
May-June 2 Apis cerana, Apis mellifera 1 S
Dahlia
daenranthema
Exotic Aug-Nov 4
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus, Bombus simillmus. Eoseristalis cerealis,
Eristalis tenax, Eristalinus aeneu, Formica, Frankliniella
sp.
7 G
Datura stramonium
L. Solanaceae Exotic May-July 3 Episyrphus balteatus 1 S
Daucus carota L. Apiaceae Exotic June-July 2 Apis cerana, Apis mellifera, Formica sp. 2 G
Delphinium roylei
Munz Ranunculaceae Native
Apis cerana, Apis mellifera, Bombus miniatus,
Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus
,
Bombus trifasciatus, Bombus simillmus,
Bombus
tunicatus
2 G
Descurainia sophia
Prantl Brassicaceae Native
May, June 2 Apis cerana, Apis mellifera, Pieris brassicae. 2 G
Digitalis purpurea
L. Scrophulariaceae
Exotic Apis cerana, Apis mellifera 1 S
Dipsacus inermis
Wall. Dipsacaceae Native
Jul-Aug 2
Apis cerana, Apis mellifera, Cetonia aurata,
Bombus
tunicatus, Bombus rufofasciatus, Bombus trifasciatus,
Bombus pyrosoma, Bombus asiaticus,
3 G
Doronicum falconeri
Hook. Asteraceae Native
Apis cerana, Apis mellifera 1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
174
Echium
plantagineum L. Boraginaceae Exotic Apis cerana, Apis mellifera 1 S
Epilobium hirsutum
L. Onagraceae Native
May, June 2 Apis cerana, Apis mellifera 1 S
Epilobium
parviflorum Schreb. Onagraceae Native
May, June 2 Apis cerana, Apis mellifera 1 S
Epilobium
royleanum Hausskn.
Onagraceae Native
May, June 2 Apis cerana, Apis mellifera 1 S
Eremurus
himalaicus Baker.
Liliaceae Native
April, May
2
Apis cerana, Apis mellifera, Episyrphus balteatus,
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale
,
Lasioglossum leucozonium
3 G
Eriobotyria
japonica
Exotic Feb-March
Oct- Dec
5
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus, Bombus simillmus, Vespa sp.
, Vanessa
cashmiriensis
3 G
Erysimum
hieraciifolium L. Brassicaceae Exotic
Apis cerana, Apis mellifera, Lasioglossum himalayense,
Lasioglossum marginatum 2 G
Eschscholzia
californica Cham. Papaveraceae Exotic April, May,
June 3
Apis cerana, Apis mellifera, Eristalis tenax, Eristalinus
taeniops 3 G
Euonymus
hamiltonianus Wall.
Celastraceae Exotic Apis cerana, Apis mellifera 1 S
Euphorbia
helioscopia L. Euphorbiaceae Exotic April-June 3 Apis cerana, Apis mellifera 1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
175
Euphorbia wallichii
Hook.f. Euphorbiaceae Native
April-June 3 Apis cerana, Apis mellifera 1 S
Foeniculum vulgare
Mill. Apiaceae Exotic June, July 2 Apis cerana, Apis mellifera, Vespa velutina, Formica sp
.,
Eristalis tenax, Eoseristalis cerealis. 5 G
Forsythia
viridissima Lindl. Oleraceae Exotic March 1
Apis cerana, Apis mellifera, Episyrphus balteatus,
Xylocopa valga, Xylocopa violacea, Pyrrhocorus,
Megachile rotundata, Halictus propinquus,
Halictus
constrictus, Scathophaga stercoraria.
7 G
Fragaria ananassa
Duchesne Rosaceae Exotic April, May
2
Apis cerana, Apis mellifera, Eristalis tenax, Eoseristalis
cerealis, Formica sp. 4 G
Fragaria nubicola
Lacaita Rosaceae Native
April, May
2
Apis cerana, Apis mellifera, Eoseristalis cerealis,
Eristalis tenax, Episyrphus balteatus, Formica. 5 G
Fumaria indica
Pugsley Papaveraceae Native
April, May
2 Apis cerana, Apis mellifera 1 S
Galinsoga
parviflora Cav. Asteraceae Exotic June-Aug 3 Episyrphus balteatus, Frankliniella sp. 2 G
Gentiana
cachemirica Decne. Gentianaceae Native
May-June 2
Apis cerana, Apis mellifera, Bombus miniatus,
Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus
,
Bombus trifasciatus, Bombus simillmus,
Bombus
tunicatus
2 G
Geranium nepalense
Sweet Geraniaceae Native
May-June 2
Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium
2 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
176
Geranium pratense
L Geraniaceae Native
May-June 2
Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium
2 G
Geranium
wallichianum Don Geraniaceae Native
May-June 2
Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium
2 G
Geum roylei Wall. Rosaceae Native
Apis cerana, Apis mellifera 1 S
Gladiolus
hortulanus Bailey Iridaceae Exotic July-Oct 4 Apis cerana,
Apis mellifera, Bombus tunicatus, Bombus
trifasciatus, Bombus simillmus. 2 G
Hackelia uncinata
Fisch. Boraginaceae Native
March- May
3 Apis cerana, Apis mellifera 1 S
Halianthus annus L.
Asteraceae Exotic July-Oct 4 Apis cerana, Apis mellifera, Xylocopa valga, Xylocopa
violacea, 2 G
Hibiscus rosa-
sinensis L. Malvaceae Exotic July-Oct 4 Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus. 2 G
Hypericum
hookerianum Wight
and Arn.
Hypericaceae Native
May, June 2 Apis cerana, Apis mellifera 1 S
Hypericum
perforatum L. Hypericaceae Native
May, June 2 Apis cerana, Apis mellifera, Eristalinus taeniops, 2 G
Iberis amara L. Brassicaceae Exotic April, May
2
Apis cerana, Apis mellifera, Calliphora lata, Eristalis
tenax, Eristalinus taeniops, 4 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
177
Impatiens
glandulifera Royle Balsaminaceae Native
Apis cerana, Apis mellifera, Bombus trifasciatus, 2 G
Indigofera
heterantha Wall Fabaceae Native
May 1 Apis cerana, Apis mellifera 1 S
Inula royleana Clark
Asteraceae Native
Apis cerana, Apis mellifera, Eristalis tenax, Vanessa
cashmiriensis. 3 G
Inula racemosa
Hook. Asteraceae Native
Apis cerana, Apis mellifera, Eristalis tenax, Vanessa
cashmiriensis. 3 G
Ipomoea tricolor
Cav. Convolvulaceae Exotic Apis cerana, Apis mellifera 1 S
Iris decora Wall.
Iridaceae Exotic May, June 2
Apis cerana, Apis mellifera, Musca domestica, Andrena,
Lasioglossum margenatum, Xylocopa valga, Xylocopa
violacea.
5 G
Iris hookeriana
Foster Iridaceae Native
March 1
Apis cerana, Apis mellifera, Musca domestica, Andrena,
Lasioglossum margenatum, Xylocopa valga, Xylocopa
violacea.
5 G
Lactuca
dolichophylla
Kitam.
Asteraceae Native
April, May
2 Apis cerana, Apis mellifera 1 S
Lagotis cashmeriana
Rupr. Scrophulariaceae
Native
Apis cerana, Apis mellifera, Macroglossum nycteris 2 G
Lamium album L. Lamiaceae Native
Apis cerana, Apis mellifera, 1 S
Lavandula
officinalis L. Lamiaceae Exotic June 1 Apis cerana, Apis mellifera, Pieris brassicae, 2 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
178
Lavatera
cashmiriana
Cambess.
Malvaceae Native
June –July 2
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus. 2 G
Leonurus cardiaca
L. Lamiaceae Native
Apis cerana, Apis mellifera 1 S
Ligularia fischeri
Turcz. Asteraceae Native
Apis cerana, Apis mellifera, Bombus tunicatus,
Episyrphus balteatus 3 G
Lindelofia longiflora
Baill.
Boraginaceae Native
Apis cerana, Apis mellifera, Mylabris pustulata,
Xylocopa
valga, Xylocopa violacea, Bombus miniatus,
Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus
,
Bombus trifasciatus, Bombus simillmus,
Bombus
tunicatus, Vanessa cashmiriensis, Episyrphus balteatus.
6 G
Lonicera japonica
Thunb. Caprifoliaceae Exotic May, June 2 Apis cerana, Apis mellifera 1 S
Lotus corniculatus
L. Fabaceae Native
May 1 Apis cerana, Apis mellifera 1 S
Lycopersicon
esculentum Mill. Solanaceae Exotic
Apis cerana, Apis mellifera, Bombus trifasciatus, Bombus
simillmus, Bombus tunicatus 2 G
Magnolia
grandiflora L. Magnoliaceae Exotic April, 1
Apis cerana, Apis mellifera, Musca domestica,
Scathophaga stercoraria. 3 G
Mahonia borealis
Takeda Berberidaceae Exotic April, May
2 Apis cerana, Apis mellifera, Eristalinus
Taeniops,
Eristalis tenax, Eristalinus taeniops, 3 G
Malus domestica
Borkh. Rosaceae Exotic April, May
2
Apis cerana, Apis mellifera, Bibio marci, Plecia
nearctica, Bombus tunicates, Andrena patella, Andrena
12 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
179
cineraria, Andrena floridula, Andrena
flavipes,
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale
,
Lasioglossum leucozonium, Episyrphus balteatus,
Megachile rotundata, Pieris brassicae, Formica,
Vanessa
cardui, Vanessa cashmiriensis, Scathophaga stercoraria.
Malva neglecta
Wallr. Malvaceae Native
May-Sept 5 Apis cerana, Apis mellifera, Ceratina hieroglyphica, 2 G
Malva sylvestris L. Malvaceae Native
May-Sept 5 Apis cerana, Apis mellifera, Ceratina hieroglyphica, 2 G
Marrubium vulgare
L Lamiaceae Native
Apis cerana, Apis mellifera 1 S
Meconopsis latifolia
Prain Papaveraceae Native
Apis cerana, Apis mellifera, Bombus trifasciatus, Bombus
simillmus, Bombus tunicatus 2 G
Medicago lupulina
L. Fabaceae Native
Apis cerana, Apis mellifera 1 S
Mentha arvensis L.
Lamiaceae Exotic June-Oct 5
Apis cerana, Apis mellifera, Scolia nobilitata, Scolia
hauseri, Scolia soror, Scoliinae proscoliinae, Eristalis
tenax, Eoseristalis cerealis
5 G
Mentha longifolia L.
Lamiaceae Native
June-Oct 5
Apis cerana, Apis mellifera, Scolia nobilitata, Eristalis
tenax, Eoseristalis cerealis. 4 G
Myosotis arvensis L.
Boraginaceae Native
Apis cerana, Apis mellifera 1 S
Myosotis caespitosa
Schultz Boraginaceae Native
Apis cerana, Apis mellifera 1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
180
Nepeta erecta
Benth. Lamiaceae Native
Apis cerana, Apis mellifera 1 S
Narcissus poeticus
L. Liliaceae Exotic March-May
3 Apis cerana, Apis mellifera, Eristalis tenax. 2 G
Nasturtium
officinale Aiton Brassicaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Nelumbo nucifera
Gaertn. Nelumbonaceae Native
July-Aug 2 Apis cerana, Apis mellifera 1 S
Nerium indicum
Mill. Apocynaceae Exotic July-Aug 2 Apis cerana, Apis mellifera 1 S
Nymphaea alba L. Nymphaeaceae Native
Apis cerana, Apis mellifera 1 S
Oxalis corniculatus
L. Oxalidaceae Native
April-May 2 Apis cerana, Apis mellifera, Pieris brassicae, 2 G
Oxytropis
cashmeriana
Cambess.
Fabaceae Native
Macroglossum nycteris 1 S
Paparver dubium L.
Papaveraceae Exotic May 1 Apis cerana, Apis mellifera 1 S
Papaver somniferum
L. Papaveraceae Exotic May 1 Apis cerana, Apis mellifera 1 S
Parthenium
hysterophorus L. Asteraceae Exotic Aug-Nov 4 Eoseristalis cerealis, Eristalis tenax, Formica 3 G
Petunia alba
Ferguson andOttley Solanaceae Exotic June-Nov 6 Apis cerana, Apis mellifera, Pieris brassicae,
Vanessa
cashmiriensis, 3 G
Phaseolus vulgaris Fabaceae Exotic May-Aug 4
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
4 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
181
L.
trifasciatus, Bombus simillmus, Xylocopa valga, Xylocopa
violacea, Andrena flavipes.
Physalis
philadelphica Lam. Solanaceae Exotic Jul-Sept 3 Apis cerana, Apis mellifera 1 S
Pinus wallichiana
A.B.Jacks. Pinaceae Native
Aug 1 Bombus tunicatus, Bombus trifasciatus, 1 S
Plantago lanceolata
L. Plantaginaceae Native
June-Aug 3 Apis cerana, Apis mellifera 1 S
Plantago major L. Plantaginaceae Native
June-Aug 3 Apis cerana, Apis mellifera 1 S
Plectranthus
rugosus Benth. Lamiaceae Native
Aug-Nov 4
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus, Bombus simillmus, Eristalinus taeniops 3 G
Poa annua L. Poaceae Native
Apis cerana, Apis mellifera 1 S
Podophylum
haxandrum Royle Berberidaceae Native
April-June 3
Apis cerana, Apis mellifera, Lassioglossum margenatum,
Lassioglossum himalayense 2 G
Polygonum
amphibium L. Polygonaceae Native
May-July 2 Apis cerana, Apis mellifera 1 S
Polygonum
amplexicaule Don Polygonaceae Native
May-July 2 Apis cerana, Apis mellifera 1 S
Potentilla
argyrophylla Lehm. Rosaceae Native
April-June 3
Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium,
2 G
Potentilla reptans L.
Rosaceae Native
April, May
2 Apis cerana, Apis mellifera, Lassioglossum nursei
,
Lassioglossum himalayense, Lassioglossum rugolatum,
2 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
182
Lassioglossum polyctor
, Lasioglossum margenatum,
Lasioglossum sublaterale, Lasioglossum leucozonium,
Prunella vulgaris L.
Lamiaceae Native
June-Sept 4 Apis cerana, Apis mellifera 1 S
Prunus armeniaca
L. Rosaceae Exotic April 1 Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Vanessa cashmiriensis, Pieris brassicae 4 G
Prunus avium L.
Rosaceae Exotic April 1
Apis cerana, Apis mellifera, Bibio marci, Plecia
nearctica, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale,
Lasioglossum leucozonium, Episyrphus
balteatus, Pieris brassicae,
Formica, Vanessa cardui,
Vanessa cashmiriensis.
8 G
Prunus domestica L.
Rosaceae Exotic April-May 2
Apis cerana, Apis mellifera, Bibio marci, Plecia
nearctica, Bombus tunicates, Scathophaga stercoraria.
Andrena patella, Andrena cineraria, Andrena floridula,
Andrena flavipes, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale,
Lasioglossum leucozonium, Episyrphus
balteatus, Megachile rotundata, Pieris brassicae,
Formica, Vanessa cardui, Vanessa cashmiriensis.
12 G
Prunus persica
Batsch Rosaceae Exotic April-May 2
Apis cerana, Apis mellifera, Bibio marci, Plecia
nearctica, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
8 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
183
sublaterale, Lasioglossum leucozonium,
Episyrphus
balteatus, Pieris brassicae,
Formica, Vanessa cardui,
Vanessa cashmiriensis.
Pseudomartensia
nemorosa Stewart
and Kazmi
Boraginaceae Native
May-July 3
Apis cerana, Apis mellifera, Bombus tunicatus, Bombus
trifasciatus. 2 G
Pteracanthus
urticifolius Bremek. Acanthaceae Native
July-Sept 3 Apis cerana, Apis mellifera 1 S
Punica granatum L.
Lythraceae Exotic May, June 2 Apis cerana, Apis mellifera, Formica, 2 G
Pyrus communis L.
Rosaceae Exotic April-May 2
Apis cerana, Apis
mellifera, Bibio marci, Plecia
nearctica, Bombus tunicates, Andrena patella, Andrena
cineraria, Andrena floridula, Andrena flavipes,
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale,
Lasioglossum leucozonium, Episyrphus balteatus,
Megachile rotundata, Pieris brassicae,
Formica, Vanessa
cardui, Vanessa cashmiriensis.
11 G
Pyrus pyrifolia
Nakai
Rosaceae Exotic April-May 2
Apis cerana,
Apis mellifera, Bibio marci, Plecia
nearctica, Bombus tunicates, Andrena patella, Andrena
cineraria, Andrena floridula, Andrena flavipes,
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale,
Lasioglossum leucozonium, Episyrphus balteatus,
11 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
184
Megachile rotundata, Pieris brassicae,
Formica, Vanessa
cardui, Vanessa cashmiriensis.
Ranunculus arvensis
L. Ranunculaceae Native
April-June 3 Apis cerana, Apis mellifera 1 S
Ranunculus laetus
Wall.
Ranunculaceae Native
April-June 3
Apis cerana, Apis mellifera, Sphecodes lasimensis,
Sphecodes tantalus, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale, Lasioglossum leucozonium,
3 G
Ranunculus
scleratus L. Ranunculaceae Native
April-June 3 Apis cerana, Apis mellifera 1 S
Raphanus sativus L.
Brassicaceae Exotic May, June 2
Apis cerana, Apis mellifera, Eristalis tenax, Pontia
daplidice, 3 G
Rhododendron
campanulatum D.
Don Ericaceae Native
May-July 3
Apis cerana, Apis mellifera, Bombus miniatus,
Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus,
Bombus trifasciatus, Bombus simillmus,
Bombus
tunicatus
2 G
Robinia
pseudoacacia L Fabaceae Exotic May 1 Apis cerana, Apis mellifera 1 S
Rorippa islandica
Borbas Brassicaceae Native
May 1 Apis cerana, Apis mellifera, Pieris brassicae 2 G
Rosa brunonii Lindl.
Rosaceae Native
May-July 3
Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Andrena flavipes, Cetonia aurata, Eristalis
tenax, Eoseristalis cerealis, Lassioglossum nursei,
11 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
185
Lassioglossum himalayense, Metasyrphus bucculatus,
Episyrphus balteatus, Scath
ophaga stercoraria,
Eristalinus taeniops
Rosa webbiana L.
Rosaceae Native
May-July 3
Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Andrena flavipes,
Cetonia aurata, Eristalis
tenax, Eoseristalis cerealis, Lassioglossum nursei,
Lassioglossum himalayense, Metasyrphus bucculatus,
Episyrphus balteatus, Scathophaga stercoraria,
Eristalinus taeniops.
9 G
Rosa indica L. Rosaceae Exotic May-Dec 7 Apis cerana, Apis mellifera 1 S
Rubus elipticus Sm. Rosaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Rubus fruticosus L. Rosaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Rubus niveus Thunb.
Rosaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Rubus occidentalis
L. Rosaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Rubus ulmifolius
Schott Rosaceae Native
May-July 3 Apis cerana, Apis mellifera 1 S
Rudbeckia hirta L. Asteraceae Exotic July-Nov 5 Apis cerana, Apis mellifera 1 S
Rumex acetosa L. Polygonaceae Native
May-June 2 Apis cerana, Apis mellifera 1 S
Salix alba L. Salicaceae Exotic April-May 2
Apis cerana, Apis mellifera, Eristalis tenax, Episyrphus
balteatus. Lasioglossum margenatum. 3 G
Salix caprea L. Salicaceae Exotic Feb., March
2 Apis cerana, Apis mellifera 1 S
Salvia hians Benth. Lamiaceae Native
May, June 2 Bombus miniatus, Bombus pyrosoma,
Bombus asiaticus,
Bombus rufofasciatus,
Bombus trifasciatus, Bombus
2 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
186
simillmus, Bombus tunicatus
Salvia moocroftiana
Benth. Lamiaceae Native
May, June 2 Apis cerana,
Apis mellifera, Bombus trifasciatus, Bombus
simillmus, Bombus tunicatus, Lasioglossum himalayense. 3 G
Sambucus wightiana
Wight and Arn. Caprifoliaceae Native
June-July 2
Apis cerana, Apis mellifera, Lasioglossum margenatum.
Formica sp., Musca domestica, Mordella sp.,
Metasyrphus bucculatus
6 G
Sanvitalia
procumbens Lam. Asteraceae Exotic Aug-Nov 4
Apis cerana, Apis mellifera, Eristalis tenax,
Sphaerophoria bengalensis, 3 G
Saussurea costus
Lipsch. Asteraceae Native
June-Aug 3
Apis cerana, Apis mellifera, Bombus miniatus,
Bombus
pyrosoma, Bombus asiaticus, Bombus rufofasciatus
,
Bombus trifasciatus, Bombus simillmus,
Bombus
tunicatus
2 G
Senecio
chrysanthemoides
DC.
Asteraceae Exotic Apis cerana, Apis mellifera 1 S
Sisymbrium irio L. Brassicaceae Exotic April, May
2 Apis cerana, Apis mellifera, Eristalis tenax 2 G
Sium latijugum
Clarke Apiaceae Native
Apis cerana, Apis mellifera, Eristalis tenax, Episyrphus
balteatus, Formica, Sphaerophoria bengalensis,
Sphaerophoria macrogaster, Sphaerophoria fatarum
5 G
Skimmia anquetilia
Shaw Rutaceae Native
May-July 3 Apis cerana, Apis mellifera, Andrena flavipes,Vespa sp.,
Formica sp. 4 G
Solanum melongena
L. Solanaceae Exotic Apis cerana, Apis mellifera 1 S
Solanum nigrum L. Solanaceae Exotic May-Nov 7 Apis cerana, Apis mellifera 1 S
Chapter – 4 Characterization of generalized and specialized plant insect interactions
187
Solanum tuberosum
L. Solanaceae Exotic May-July 3 Apis cerana, Apis mellifera 1 S
Sophora japonica L.
Fabaceae Exotic May-June 2 Apis cerana, Apis mellifera 1 S
Sonchus oleraceus
L. Asteraceae Exotic May-Oct 6 Apis cerana, Apis mellifera, Frankliniella sp. 2 G
Stachys floccosa
Benth. Lamiaceae Native
June 1 Apis cerana, Apis mellifera 1 S
Stellaria media Vill.
Carophyllaceae Native
Jan-May 5
Apis cerana, Apis mellifera, Lassioglossum nursei,
Lassioglossum himalayense, Lassioglossum rugolatum,
Lassioglossum polyctor, Lasioglossum margenatum,
Lasioglossum sublaterale,
Lasioglossum leucozonium,
Episyrphus balteatus, Sphaerophoria bengalensis,
Sphaerophoria macrogaster, Eristalis tenax.
6 G
Sternbergia lutea
Spreng. Liliaceae Exotic Jan- March
3
Apis cerana, Apis mellifera, Pieris brassicae, Vanessa
cashmiriensis,
Episyrphus balteatus, Scathophaga
stercoraria.
5 G
Syringa emodi Royle
Oleaceae Native
Apis cerana, Apis mellifera 1 S
Tagetes patula L.
Asteraceae Exotic Aug-Dec 5
Apis cerana, Apis mellifera, Bombus tunicatus,
Bombus
trifasciatus, Pieris brassicae,
Vanessa cashmiriensis,
Eristalis tenax, Eoseristalis cerealis, Colias philodice,
Issoria lathonia, Hesperia comma
9 G
Taraxacum
officinale Wigg. Asteraceae Native
March-June
and Sept-
Nov
7
Apis cerana, Apis mellifera, Vespa velutina, Colias
philodice, Colias fieldi, Hesperia comma, Pieris
brassicae, Formica, Mylabris pustulata, Eristalis tenax,
12 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
188
Eoseristalis cerealis, Andrena flavipes. Calliphora lata,
Vanessa cardui, Lassioglossum nursei, La
ssioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum, Lasioglossum
sublaterale, Lasioglossum leucozonium,
Thymus lineasis
Benth. Lamiaceae Native
April, May
2 Apis cerana, Apis mellifera Vanessa
cashmiriensis,
Eristalis tenax. Polyommatus icarus, Danais chrysippus. 5 G
Thymus serphyllum
L. Lamiaceae Native
April, May
2
Apis cerana, Apis mellifera, Vanessa cashmiriensis,
Eristalis tenax, Polyommatus icarus, Danais chrysippus. 5 G
Trifolium pratense
L.
Fabaceae Native
April-Nov 8
Apis cerana, Apis mellifera, Bombus tunicatus,
Bombus
trifasciatus, Pieris brassicae,
Vanessa cashmiriensis,
Vanessa cardui,
Hesperia comma, Polyommatus icarus,
Andrena flavipes, Issoria lathonia, Pontia daplidice,
Lassioglossum nursei, Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale,
Lasioglossum leucozonium,
10 G
Trifolium repens L.
Fabaceae Native
April, May,
June 8
Apis cerana, Apis mellifera, Bombus tunicatus,
Bombus
trifasciatus, Pieris brassicae,
Vanessa cashmiriensis,
Vanessa cardui,
Hesperia comma, Polyommatus icarus,
Andrena flavipes, Issoria lathonia, Pontia daplidice
.Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale,
Lasioglossum leucozonium,
10 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
189
Tulipa stellata
Hook.
Liliaceae Native
April,May 2
Apis cerana, Apis mellifera,
Lassioglossum nursei,
Lassioglossum nursei,
Lassioglossum himalayense,
Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum margenatum, Lasioglossum sublaterale,
Lasioglossum leucozonium,
2 G
Valeriana
hardwickii Wall. Caprifoliaceae Native
April Apis cerana, Apis mellifera 1 S
Verbascum thapsus
L. Scrophulariaceae
Native
June Apis cerana, Apis mellifera, Formica 2 G
Veronica arvensis L.
Scrophulariaceae
Native
Jan- May
and Nov-
Dec
7
Apis cerana, Apis mellifera,
Andrena floridula, Pieris
brassicae,
Vanessa cashmiriensis, Scathophaga
stercoraria, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale, Lasioglossum leucozonium,
6 G
Veronica persica
Poir.
Scrophulariaceae
Native
Jan- May
and Nov-
Dec
7
Apis cerana, Apis mellifera,
Andrena floridula, Pieris
brassicae, Vanessa
cashmiriensis, Scathophaga
stercoraria, Lassioglossum nursei,
Lassioglossum
himalayense, Lassioglossum rugolatum,
Lassioglossum
polyctor, Lasioglossum margenatum,
Lasioglossum
sublaterale, Lasioglossum leucozonium,
6 G
Viburnum
grandiflorum Wall.
ex DC.
Adoxaceae Native
Jan- May 4
Apis cerana, Apis mellifera, Pieris brassicae, Vanessa
cashmiriensis, Vanessa cardui, Eristalis tenax,
Pyrrhocorus, Colias philodice, Colias fieldi.
6 G
Viola biflora L. Violaceae Native
May 1 Apis mellifera, Apis cerana, Pieris brassicae,
Vanessa
3 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
190
cashmiriensis
Viola odorata L. Violaceae Native
March,
April 2
Pieris brassicae, Vanessa cashmiriensis, Halictus
constrictus, 3 G
Weigela floribunda
C.A.Mey. Caprifoliaceae Exotic April-May 2 Apis cerana, Apis mellifera 1 S
Zea mays L. Poaceae Exotic
Apis cerana, Apis mellifera, 1 S
Zinnia elegans L.
Asteraceae Exotic Aug-Dec 5
Apis cerana, Apis mellifera, Xylocopa valga,
Xylocopa
violacea, Bombus tunicatus,
Bombus trifasciatus, Bombus
simillmus, Eristalis tenax, Am
egilla fallax, Pieris
brassicae, Vanessa cashmiriensis,
Hesperia comma,
Eoseristalis cerealis, Issoria lathonia, Frankliniella sp.
11 G
Chapter – 4 Characterization of generalized and specialized plant insect interactions
191
DISCUSSION
The Himalayas represent one of the world’s highest ranges of
mountains, which are located in the Indian subcontinent. Being one of the
global biodiversity hotspots, the Himalayas harbour a precious biological
wealth of great economic value and scientific interest. Lying in its north-
western part is the Kashmir Himalaya, which is renowned for its natural
landscape characterized by a diverse and endemic biodiversity (Lawrence
1895; Dar et al., 2013). Notwithstanding rich biodiversity, very few research
studies have been carried out to explore the diversity of flowering plants and
their insect visitors in this Himalayan region.
The present results revealed that in Kashmir Himalaya, the plant
pollinator web is the mixed interaction of four different interacting pillars such
as generalized plants, specialized plants, generalized pollinators and specialized
pollinators shaping four different types of plant pollinator interactions (Fig.
4.8). The plant pollination web of Kashmir Himalaya consists of 215 flowering
plant species which are interacting with 70 flower visitor species (Table 4) is
in accordance with Paray et al. (2014), Ganie et al. (2013), Gomez and Zamora
(1999). Out of 215 plant species majority (134) are generalist serving pollen
and nectar for a large number of pollinators (Kandori 2002; Memmott, 1999)
and generalization in plant pollinator interaction is predominant and most
diverse group of pollinators are associated with it (Herrera, 1996; Waser et al.,
1996; Olesen, 2000). A single plant pollinator web is formed by different plant
pollinator interactions in a cumulative manner (Memmott, 1999), in which
different pollinator species are interacting with different foraging plant species
and vice versa reflecting both generalization as well as specialization
interactions of plant pollinator web.
Out of 215 flowering plant species 121 plant species are native whereas
94 plant species are exotic to Kashmir Himalaya (Fig. 4.6). Both types of plant
species are the part and parcel of plant pollinator web existing in present study
Chapter – 4 Characterization of generalized and specialized plant insect interactions
192
area. Among 70 species of flower visitors, Apis mellifera is exotic to this
region interacting with 204 flowering plants both native as well as exotic
specifying its role as generalist pollinator competting our indigenous Apis
cerana which interacts with 205 flowering plant species (Fig. 4.2). Our result
revealed that flowering plant species blooming after the onset of winter act as
generalists for flower visitors as compared to later flowering and those which
has long flowering period as compared to short period flowering period.Among
flowering plant species Malus domestica and Prunus domestica are at the top
of generalization specialization continuum interacting with 23 flower visitor
species followed by Pyrus communis and Pyrus pyrifolia (22) Trifolium
pratense and Trifolium repense are foraged by 21 species of flower visitors
(Table 4.1). Bird as a flower visitor was not found and is not included in the
plant pollinator web of Kashmir Himalaya (Cronk and Ojeda, 2008).
Our results revealed that generalist plant species interacts with
generalist as well as specialist pollinator species and specialist plant species
interacts with generalist or specialist pollinator species (Table 4.1) is in
accordance with Herrera 2005. Our result revealed that there is 62%
generalization in plant pollinator interaction and 38% specialization, a need of
concern due to a good percentage of interaction is specialized which exists
(Pellmyr, 2002) and prone to adverse effect of pollinator decline. Specialized
plant pollinator interaction should be focussed and studied in depth for
conservation purpose in order to keep ecosystemic outcome sustainable.
Sustenance of 75% of crop plants species, 35% of crops production (Klein,
2007) and 90% flowering plants (Ollerton et al., 2011) are dependent on plant
pollinator web. So it becomes necessary to study the ecology of plants and their
flower visitors and the structure of plant pollinator web (Corbet, 2000).
Understanding the structure of plant pollinator web of Kashmir Himalaya shall
help us to generate income as well as contribute food production by enhancing
pollination in pollinator dependent crops (Bailes et al., 2015). Our results
Chapter – 4 Characterization of generalized and specialized plant insect interactions
193
revealed that the present plant pollinator web has its ecological as well as
evolutionary significance. Introduction of exotic pollinator (Apis mellifera) or
foraging plant (Abelia grandiflora) has both positive as well as negative
consequences and may produce competition or facilitation among pollinators or
foraging plant species (Thomson, 2006; Stout and Morales, 2009; Kearns and
Inouye1997; Carolina et a.,l 2006). Competition among pollinator pool is
enhanced by introducing an exotic pollinator species (Potts et al., 2010). Out of
215 plant species 204 plants are foraged by the exotic pollinator species (Apis
mellifera) with almost very wide range of feeding sites as compared to native
pollinator species (Megachile rotundata, Eristalis tenax etc.). Some native
pollinator species have very narrow range of feeding viz. Spilomyia sulphurea,
Ceratina propinqua etc. and are prone to decline. Co-existence of exotic and
native pollinators in a web generates competition which produces negative
impacts on both exotic as well as native pollinator (Thomson 2006; Goulson
and Sparrow, 2009), especially native specialists (Traveset and Richardson,
2006) as compared to native generalists (Potts et al., 2010). Introduction of
honey bee (Apis mellifera) to various non local areas developed competition for
feeding resource (pollen and nectar) with the endemic bee fauna, pushing them
to expand the boundary of their activity (Kearns and Inouye, 1997). Honey
bees were found to dominate the flower mate as compared to native bees
(Ginsburg, 1983)
The generalist plant species viz. Malus domestica and Prunus
domestica which are exotic in origin harbour the maximum number of
pollinator taxa as compared to indigenous plant species viz. Viburnum
grandiflorum (Table 4.1) and may affect the indigenous flower-pollinator webs
by outcompeting the native foraging plants by their floral advertisement and
reward consequently reducing the reproductive success of native plants
(Dietzsch et al., 2011) as native pollinators prefer exotic flowering plants for
food purposes leaving native plants as such (Waring et al., 1993) and on the
Chapter – 4 Characterization of generalized and specialized plant insect interactions
194
other side exotic plants (Eriobotyria japonica) act as best feeding sources for
native pollinators under food scarity condition e.g. exotic plant Impatiens
glandulifera acts as feeding site for bumble bees (Kleijn and Raemakers,
2008).Decrease in the resource diversity for pollinators eliminates the
specialized pollinators and affects the architecture of flower pollinator
interaction (Burkle 2013, Williams and Osborne, 2009). Plant pollinator web of
Kashmir Himalaya is different from the general web as more specialization in
plant species is found relying on small pollinator taxa and structure and
composition of this web may change either towards specialization or
generalization with respect to place and time. Our results predict that the plant
pollinator web may be affected by the climate change (Hegland et al., 2008).
Thus to acquire plant pollinator interaction knowledge, it is compulsory
for understanding the structure and fuction of an ecosystem and developing
guidlines for management and conservation of ecosystem.
CONCLUSIONS
The present study revealed that 215 plant pollinator interactions 134
(62%) are generalized whereas 81 (38%) are specialized. These two sets of
interactions involve 215 foraging plant species and 70 different flower visitor
species. Among 215 different flowering plant species 03 plant species are
foraged by 01 flower visitor species, 79 plant species foraged by 2 species of
flower visitors, 22 plant species by 3 flower visitor species, 20 plant species by
4 flower visitor species, 11 plant species by 5 flower visitor species, 11 plant
species by 6 flower visitor species etc. Plant generalization specialization
continuum revealed that Malus domestica and Prunus domestica are foraged by
the maximum number (23) of flower visitor species belonging to 12 genera
followed by Taraxacum officinale are foraged by 21 species belonging to 12
Chapter – 4 Characterization of generalized and specialized plant insect interactions
195
genera of insect visitors. Pyrus communis and Pyrus pyrifolia are foraged by
22 species belonging to 11 genera of flower visitors. Pinus wallichiana was
foraged by Bombus tunicatus and Bombus trifasciatus, Aster thomsonii was
foraged by Lassioglossum margenatum, Lassioglossum himalayense, and
Galinsoga parviflora by Episyrphus balteatus, Frankliniella sp. Oxytropis
cashmeriana is foraged by Macroglossum nycteris only, Datura stramonium
and Bidens tripartita by Episyrphus balteatus only. 75 plant species are
foraged only by Apis cerana and A. mellifera.
Apis cerana is the most generalist pollinator visitor foraging 205 plant
species followed by Apis mellifera foraging 204 plant species Eristalis tenax
(41), Bombus trifasciatus (38), Bombus tunicatus (37), Lasioglossum
margenatum (35), Lassioglossum himalayense (30), Lassioglossum nursei (29),
and Episyrphus balteatus, Lassioglossum rugolatum, Lassioglossum polyctor,
Lasioglossum sublaterale, Lasioglossum leucozonium by 26, Pieris brassicae
forages 25 plant species, Xylocopa valga, Xylocopa violacea forages 24 plant
species, Formica sp., Andrena flavipes forages 22 plant species, Vanessa
cashmiriensis forages 21 plant species, Bombus simillmus, Eoseristalis cerealis
forages 19 plant species, Vanessa cardui forages 14 plant species, Scathophaga
stercoraria forages 13 plant species, Bombus rufofasciatus, Bombus pyrosoma
forages 12 plant species, Bombus asiaticus, Eristalinus taeniops and
Megachile rotundata forages 10 plant species, Bombus miniatus forages 9
plant species, Andrena patella, Andrena floridula forages 8 plant species,
Ceratina hieroglyphica, Hesperia comma forages 7 plant species, Frankliniella
sp., Sphaerophoria bengalensis, Bibio marci, Plecia nearctica, Cetonia aurata
forages 6 plant species, Eristalinus aeneu, Metasyrphus bucculatus, Musca
domestica, Polyommatus icarus, Danais chrysippus forages 5 plant species,
Sphaerophoria macrogaster, Issoria lathonia forages 4 plant species, Mordella
sp., Amegilla fallax, Vespa velutina, Pyrrhocorus sp., Colias philodice, Pontia
daplidice forages 3 plant species, Ceratina propinqua, Sphecodes lasimensis,
Chapter – 4 Characterization of generalized and specialized plant insect interactions
196
Sphecodes tantalus, Coccinella magnifica, Mylabris pustulata, Scolia
nobilitata, Halictus constrictus, Colias fieldi, Macroglossum nycteris forages 2
plant species, Macroglossum nycteris, Heriades sp., Scolia hauseri, Scolia
soror, Scoliinae proscoliinae, Spilomyia sulphurea forages only one plant
species. The present study revealed that 130 interactions are found to exist
between generalized plant species and generalized flower visitor species. 82
interactions are found to exist between specialized plant and generalized flower
visitor. 6 interactions are found to exist between generalized plant species and
specialized flower visitor and non interaction was found existing between
specialized plant species and specialized flower visitor species.
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a
Plate
1.1 : Collection of insect
visitors
method. (e) insect box. (f)
c
e
a
b
visitors
(a & b) Killing jar method and (c
&
Insect cabinet.
c
d
f
e
a
e
Plate 1.2: (a) Apis cerana, (b)
Bombus
grandiflora. (d) Apis mellifera
c
e
a
b
f
Bombus
trifasciatus and (c) Vespa sp. on
Campsis
mellifera group on Cucurbita maxima. (e)
c
d
e
f
a
Plate
1.3: (a) Pyrus communis
flower
mellifera on Prunus domestica
c
e
b
flower
as mating site for Bibio sp.
(b)
domestica. (c) Xylocopa valga and (d)
d
f
a
Plate 1.4: (a) Apis mellifera on
Sonchus
Thymus serphyllum.
Sternbergia lutea. (d) Apis
c
e
b
Sonchus
oleraceus. (b) Mylabris
pustulata
(c) Apis cerana nectarivorous
activity
Apis cerana and (e) Apis mellifera on
d
f
a
c
e
Plate
1.5: Xylocopa valga (a)
before
canadensis.(c) Apis
mellifera
c
e
a
b
d
f
before
landing and (b) after landing
mellifera
on Cucumis sativus. (d) Apis
mellifera
c
d
e
f
Plate 1.6: (a) Andrena cineraria.
(d) Bombus
trifasciatus
cashmiriensis.
a b
(b) Andrena Patella. (c) Bombus
tunicatus
trifasciatus
. (e) Bombus simillmus. (f)
c d
e f
a
Plate
1.7: (a) Xylocopa valga.
(b)
c
e
a
b
(b)
Xylocopa violacea. (c)
Lassioglossum
c
d
e
f
a
Plate 2.1: (a) Bombus tunicatus on Abelia
e
c
e
a
b
Abelia grandiflora. (b) Apis mellifera on Ageratum
c
d
e
f
a
e
Plate
2.2 : (a ) Apis mellifera on
Hypericum perforatum. (c)
c
e
a
b
f
Hypericum hookerianum.(b) Apis
mellifera
(c) Xylocopa violacea on Iris decora. (d)
d
c
f
e
a
e
Plate
2.3: (a) Apis cerana
Pollenivorous
nectarivorous activity
on
indica. (d) Apis cerana
on
on Plectranthus rogosus.
c
e
a
b
f
Pollenivorous
activity and b) Apis
mellifera
on
Kniphofia uvaria.(c) Apis mellifera
on
Verbascum thapsus. (e) Bombus
tunicatus
(e) Apis mellifera on Trifolium pratense
c
d
e
f
a
d
Plate 3.1: Pollen deposition on
stigmas
rosa-sinensis. (c)
Sonchus
Colchicum luteum.
(f)
c
e
a
b
stigmas
of (a) Cichorium intybus.(b)
Sonchus
oleraceus.(d) Taraxacum
officinale
(f)
Punica granatum.
c
d
e
f
a
Plate 3.2: Pollen deposition
on
Epilobium
hirsutum
cashemiriana.(e) Emasculated
c
e
b
a
on
stigmas of (a) Punica
granatum
hirsutum
. (c)Rosa indica. (d)
Lavatera
Emasculated flower and (f) pollen loaded
c
d
f
e
a
Plate 4.1: (a) Apis cerana, (b)
Bombus
of Bombus tunicates
comma depicts
mating
simillimus on Dahlia daenranthema
c
e
a
b
Bombus
tunicatus, (c) high pollinator
along mature flower row, (d)
Hesperia
mating
site, (e) Megachile sp.,( f)
Bombus
daenranthema.
c
d
e
f
a
e
Plate
4.2: (a) Apis cerana and (b)
Eristalinus
view (d) Side view of
Apis
brassicae on Buddleja davidii
c
e
a
b
f
Eristalinus
sp. on Berberis lycium. (c)
Apis
mellifera on Brassica compestris.
(e)
davidii. (f) Eristalis tenax on Sisymbrium
c
d
e
f
a
Plate
4.3 : (a) Bombus
trifasciatus
pratense.(c) Apis cerana
c
e
a
b
trifasciatus
and (b) Bombus tunicatus on
Trifolium
cerana and (d) Sphaerophoria bengalensis
c
d
e
f
a
Plate 4.4 : (a) Apis mellifera
on
lathonia, (d)
Bombus
c
e
a
b
on
Viola biflora.(b) Apis cerana, (c)
Bombus
trifasciatus, (e) Bombus tunicatus
on
c
d
f
e
a
Plate 4.5: (a) Apis mellifera
approaching
mellifera on
Cucumis
maxima. (e) Apis
mellifera on Zinnia
c
e
a
b
approaching
to and (b) on Salix caprea
.
Cucumis
sativus. (d) Apis melliferas on
Cucurbita
mellifera on Actinidia deliciosa.
elegans.
c
d
e
f
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