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BEES: AN INTRODUCTION
Bees are found in all ecosystems on earth that support some angiosperms, since the requirement for pollen and nectar is absolute for
the development and survival of these hymenopteran insects. As such, barring the frigid cold Antarctic, bees may be found wherever there
are entomophilous flowers and the reliance of bees on flowers has, in time, resulted in the evolution of strong mutualism between the
flowering plants and their pollinator bees. It is hypothesized that bees originated from predatory wasps (of the family Sphecidae)
approximately 130 million years ago, shortly after the evolution of the first flowers, in the Cretaceous (Gupta 2014). The first bees were
solitary creatures just like the Sphecid wasps and nested in cavities in woods or burrows in soil, evolving into eusocial colonial forms
approximately 80 million years ago. The complex life histories of these eusocial bee species, involving coexistence of overwintering
broods and adults along with eggs within a single hive, necessitated provisioning the nest with food even when external sources of nutrition
dwindled, leading to the behaviour of making and storing honey from collected nectar. Evolution thus resulted in the emergence of honey-
making hive-nesting bees, all grouped under one genus Apis. The continent of Asia, chiefly its southern parts, has been suggested as the
cradle for the origin and diversification of the honeybees (Gupta 2014).
HONEYBEES: DIVERSITY
Bees belonging to the tribe Apini under the subfamily Apinae and the family Apidae are regarded as the true honeybees, of the genus
Apis, and have long been commercially treasured for viable economic returns. The genus Apis is a small and morphologically and
behaviourally unified group (Michener 2007) characterized by hairy eyes and comb building activities along with distinct division of
labour among the different castes (typically a fertile female – the queen, a few fertile males – the drones, and majority of sterile females –
the workers) that constitute each colony. This genus of bees has a wide distribution occurring in the Palearctic region (north to southern
Norway) and the Pacific maritime provinces of Russia as well as in the entirety of the African and Oriental regions (Michener 2007).
INDIGENOUS AND INTRODUCED HONEYBEES OF INDIA:
BRIEF OVERVIEW OF INTERACTION
1 1 2
Manjishtha Bhattacharyya *, Susanta Kumar Chakraborty and Sankar Kr. Acharya
1Department of Zoology, Vidyasagar University, Midnapore 721102
Email: *manjishtha.b@gmail.com, susantachakraborty@yahoo.com
2Department of Agricultural Extension, Bidhan Chandra Krishi Viswavidyalay, Mohanpur, Nadia 741252.
Email: acharya09sankar@gmail.com
*Corresponding author
Vol. No. 23(2) : March, 2017
6ENVIS Newsletter
Additionally, human activities have further extended the distribution of certain species of Apis bees, and now these bees occur in regions
where there were no previous records of such insects, notably in Australia and New Guinea.
Following the naming of the genus by Linnaeus in 1758, several species and subspecies of this genus have been described by different
authors from all over the world, and at present at least nine different species are recognized under this one genus Apis which have been
informally classified into three groups on the basis of their phylogeny (Alexander 1991; Engel 1999). The group consisting of the small
Apis bees, A. florea Fabricius and A. andreniformes Smith, constitute the basal branch of the phylogenetic tree. The large species A. dorsata
Fabricius, A. laboriosa Smith, A. binghami Cockerell and A. breviligula Maa, constitute the group of giant Apis bees which are the next
branch in the phylogenetic tree, although the last two species in this group are regarded as allopatric species of A.dorsata, and not true
species, by Michener (2007). Finally, the last and the most recent group of Apis bees in the phylogenetic tree constitute of the medium sized
bees A. mellifera Linnaeus, A. cerana Fabricius, A. koschenikovi Buttel-Reepen, A. nigrocincta Smith and A. nuluensis Tinger, Koeniger,
and Koeniger. Of all these species of honeybees, only Apis mellifera is non-native to Asia, although there have been suggestions of a
possible Asian origin of the early lineage of honeybees that gave rise to the present day European honeybee A.mellifera (Gupta 2014).
HONEYBEES: THE INDIAN SCENARIO
At present, in India, five species of naturally occurring honeybees share the diverse floral resources with one species of introduced
honeybee. All these species of bees, by virtue of being polylectic in their food preference, serve as important pollinators of the cultivated
crops and wild plants of India, apart from being providers of valuable honey and beeswax. A brief description of these honeybees is given
below.
A. Indigenous species of honeybees in India
1. Apis dorsata: Commonly known as the Giant or Rock bee, this is a large species of honeybee, with worker length of 16-18 mm.
These bees, build single exposed combs on rock walls or suspended from tree branches, and are found throughout India. Combs may often
be more than a meter in width and have distinct storage parts including honey storage at the top, followed by pollen storage, worker brood
and drone brood. Communications among workers via bee dances occur on the vertical face of the comb. These bees are highly aggressive,
although, due to large yield of honey (>30 kg per comb per year), their combs are frequently pillaged by locals. They are also highly
migratory and have strong swarming tendencies.
2. Apis laboriosa: Commonly known as the Himalayan cliff bee, this is the world's largest species of honeybee with workers
measuring as much as 3 cm in length. These build single exposed combs in high altitudes (2500m-3500m) usually hanging from vertical
cliff overhangs. Their red honey is highly prized for its medicinal properties, chiefly by ethnic mountainous tribes of India, Nepal, Bhutan
and Korea.
3. Apis cerana: This is the indigenous Asiatic honeybee (or the Eastern honeybee) and is a medium sized insect with workers
measuring approximately 11-12 mm. These are cavity nesting bees and usually build multiple combs parallel to one another with a definite
'bee space' maintained between two combs. The outer combs usually store honey, while the brood cells are in the inner combs. Worker
brood cells are smaller than drone brood cells, while the queen cells are irregular and built on the lower edge of the comb. The yield of
honey (approximately 6-8 kg per colony per year) is less than that of A. dorsata and A. laboriosa, but due to their docile nature, they can be
domesticated in conventional Langstroth hives, and are the popular choice for bee keeping throughout India. Three subspecies of this
honeybee are recognized in India – Apis cerana indica, Apis cerana cerana and Apis cerana himalaya.
4. Apis florea: Commonly known as the little / dwarf bee, this is the smallest species of honeybee in the world, both in terms of the size
of workers (length <8 mm) and the size of the comb it builds. A single exposed comb is built by this bee, hanging from tree branches, rock
cliffs, or even from rafters of houses. A unique and primitive feature of this bee, which it shares with its sister species A. andreniformis, is
the performance of the bee dance on the horizontal surface of the comb where the comb is suspended from a support (viz., tree branch).
These bees are more tolerant to higher temperatures and yield small quantities (< 0.5 – 2 kg per comb per year) of usually unifloral honey.
These can be found all over the country.
5. Apis andreniformis: These bees were previously considered as a subspecies of A. florea. The abdomens of workers in this species
are darker than the abdomens of A. florea, which is one of the key identifying features of this species. This species has been typically
observed in high elevations and in tropical forests with reduced sunlight penetration.
B. Introduced species of honeybee in India
Apis mellifera: The European honeybee was a native to Africa, most of Europe and the Middle East, and was subsequently introduced
by man to the Americas, Australasia and much of the rest of the world. Besides being managed extensively for commercial beekeeping in
India, escaped colonies of this introduced bee has populated the wilds and co-exists with its Asian counterpart A.cerana. A. mellifera is
bigger than A. cerana, but its habits in the wild are same as the latter species. The yield of honey (25-40 kg per colony per year) is higher
than the Asian bee and they are much more docile and less prone to absconding than A. cerana.
7
INTERACTIONS BETWEEN INDIGENOUS AND INTRODUCED HONEYBEE SPECIES
The different species of Apis bees occurring sympatrically have been shown to display a myriad of interactions including competition,
interference, parasitism, avoidance etc. among each other (Koeniger 1982). A brief summary of the recorded interactions between the
different species of honeybees native and introduced to Asia, including India, is listed in table 1.
It has been suggested that interspecific interactions among the different species of honeybees, endemic to Asia, have shaped their
evolution (Yang, 2009). Through fine partitioning of niches, temporally (as has been reported by Oldroyd, Rinderer and Wongsiri 1992,
among A. cerana, A. dorsata, A. florea and A. andreniformis with relation to foraging time over king palm Archontophoenix alexandrea, in
Thailand), or spatially, (Yang 2009), and even through delicate character displacements, viz., as seen in the variable length of glossa
(Oldroyd and Wongsiri 2006) or the differences in Excess Power Index (EPI) that determines the flight ability of honeybees (Hepburn,
Radloff and Fuchs 1999; Radloff, Hepburn and Koeniger 2003), the different Asian honeybee species have been able to survive, diversify
and coexist successfully.
INTRODUCED HONEYBEES IN INDIA: RISK OF INVASIVENESS?
Commercial beekeeping using the docile and manageable strains of the European honeybee Apis mellifera has seen a steady rise in
Asia, including India. Following its introduction in Himachal Pradesh in the 1960s, the rearing of this bee has increased manifold across
several states of the country, and at present in India, commercial migratory bee keeping relies heavily on this introduced bee species (Rao
and Rao 2011), much like the rest of the world. Since this bee is the European counterpart of the Asian honeybee Apis cerana, much of its
behaviour and requirements are same as the latter species, resulting in direct competition between the two species, usually for food, in
nature (table 1). Under experimental conditions, also, Apis mellifera have been shown to prevent Apis cerana from accessing food from
artificial feeding stations (Dhaliwal and Atwal 1970). Apis mellifera have been shown to be aggressive robbers of food stores from Apis
cerana colonies (Yang 2001) and because of a
supposed absence of pre-mating barriers
between these two species (Ruttner 1988) and
similar queen pheromones, frequent copulation
attempts by A. mellifera drones is made on A.
cerana queens, outcompeting the A. cerana
drones (Muzaffar and Ahmed 1990), and with a
noxious effect on the queen (Yang 2009). These
negative interactions between the two species
lead to the speculation of whether the European
honeybee Apis mellifera shall emerge as a
potential invasive threat to India's indigenous
honeybee species, A. cerana.
Apis cerana populations have been reported to have declined steadily following introduction of Apis mellifera bees, in India (Atwal
and Sharma 1971; Verma 1993), as well as in other Asian countries (Japan: Sakagami 1959; China: Yang 2005; Nepal: Partap 1998). An
increased interest in beekeeping with the European honeybees, due to their large yield of honey, beeswax and propolis, has further resulted
in a decline in conservation and management practices involving the Asian honeybee species. Although, not yet threatened with
extirpation, unless proactive conservation measures are adopted, India may see a severe dwindling in the population of its indigenous
honeybee species, with increasing importance given to commercially viable colonies of imported and introduced European honeybee
races.
At present, several behavioural and physiological differences between the indigenous Apis cerana and the introduced Apis mellifera
are serving as checks against replacement of the former species by the latter, by creating enough niche separation between the two bee
species. For instance, A. cerana have been found to forage earlier in the day and also tolerate lower temperatures than A. mellifera (Partap,
Shukla and Verma 2000). Apis cerana also have been reported to be efficient pollen collectors from heterogenous flower patches, spending
less time per flower, while Apis mellifera preferred relatively homogenous big flower patches and had greater flower processing times
(Partap et al. 2000). These differences in timing and flower patch preferences may be enough to avoid competitive exclusion of Apis
cerana by Apis mellifera. Besides, A. mellifera are more susceptible to Varroa mite infestation, which is a natural parasite of A. cerana and
causes the latter species much less harm.
However, increasing air temperatures, widespread use of pesticides and acaricides to control Varroa mites, and monoculture of
entomophilous crops may turn the tables and break these barriers stopping the introduced Apis mellifera from becoming invasive with the
elimination of its Asian counterpart Apis cerana.
Table-1. Interactions recorded between different honeybee species occurring in Asia
(both endemic and introduced)
Between species Interaction Type Reference
Apis florea & Apis andreniformis Avoidance Rinderer et al. 2002
Apis dorsata & Apis cerana Competition for food Koeniger and Vorwohl, 1979
Apis cerana & Apis mellifera Competition for food Partap, 1998; Sharma et al. 2000
Apis florea & Apis mellifera Nest robbing Koeniger 1976
Apis dorsata & Apis cerana Nest robbing Atwal and Dhaliwal 1969.
Apis mellifera & Apis cerana Mating Ruttner and Maul 1983
Apis mellifera & Apis florea Mating intervention Koeniger 1976
Vol. No. 23(2) : March, 2017
8ENVIS Newsletter
CONCLUDING THOUGHTS
The introduction of the European honeybee species Apis mellifera has boosted beekeeping and added to the rural economy of India.
It's contribution in migratory beekeeping is undeniable. However, because of its active interference with foraging and mating habits of the
Asian honeybee Apis cerana, the introduced European honeybee faces the risk of being tagged an invasive threat to the indigenous species
of honeybees in India. Timely corrective actions, however could mitigate the problems and ensure co-existence of the two species.
Through careful conservation practices, including preservation of wild flora and suitable nesting places, and controlled use of
agrochemicals and biocides, the populations of India's indigenous honeybees may be protected. Apiculture using Apis cerana subspecies,
if encouraged, could also aid in boosting the populations of this species across the varied landscapes of India, especially in places where,
forage is available marginally and insufficiently to sustain the introduced European species. The European bee species, because of its high
economic viability, both as a producer of a marketable product and as an important ecosystem service provider, at present in India may be
monitored and managed closely so that its impact on the indigenous apifauna is maintained at a minimum.
ACKNOWLEDGEMENT
The authors wish to thank University Grants Commission (UGC-India) for providing financial assistance in the form of Junior
Research Fellowship to the first author during the preparation of this manuscript.
REFERENCES
Alexander BA (1991). Phylogenetic analysis of the genus Apis. Annals of the Entomological Society of America. 84: 137–149.
Atwal AS, Dhaliwal GS (1969). Robbing between Apis indica F. and Apis mellifera L. American Bee Journal. 109: 462–463.
Atwal AS, Sharma OP (1971). The dominance of Apis mellifera over Apis indica. American Bee Journal 111(9): 343.
Dhaliwal GS, Atwal AS (1970). Interspecific relations between Apis cerana indica and Apis mellifera. Journal of Apicultural Research. 9:
53–59.
Engel MS (1999). The taxonomy of recent and fossil honeybees. Journal of Hymenoptera Research. 8: 165–196.
Gupta RK (2014). Taxonomy and distribution of different honeybee species. In Gupta RK, Reybroeck W, van Veen JW, Gupta A (Eds)
Beekeeping for Poverty Alleviation and Livelihood Security: Vol. 1: Technological Aspects of Beekeeping. 63–103. Published by
Springer Science, Netherlands.
Hepburn HR, Radloff SE, Fuchs S (1999). Flight machinery dimensions of honeybees, Apis mellifera. Journal of Comparative Physiology
B 169: 107–112.
Koeniger N (1976). Interspecific competition between Apis florea and Apis mellifera in the tropics. Bee World 57: 110–112.
Koeniger N (1982). Interactions Among the Four Species of Apis. Published by Westview Press, Boulder, Colorado, USA.
Koeniger N, Vorwohl G (1979). Competition for food among four sympatric species of Apini in Sri Lanka: Apis dorsata, Apis cerana, Apis
florea and Trigona iridipenis. Journal of Apicultural Research 18: 95–109.
Michener CD (2007). The Bees of the World (2nd Edition). Published by Johns Hopkins University Press, Baltimore, Maryland, USA.
Muzaffar N, Ahmad R (1990). Apis spp. (Hymenoptera, Apidae) and their distribution in Pakistan. Pakistan Journal of Agricultural
Research 11(1): 65–69.
Oldroyd BP, Rinderer TE, Wongsiri S (1992). Pollen resource partitioning by Apis dorsata, A. cerana, A. andreniformis, and A. florea in
Thailand. Journal of Apicultural Research 31: 3–7.
Oldroyd BP, Wongsiri S (2006). Asian Honey Bees: Biology, Conservation and Human Interactions. Published by Harvard University
Press, Massachusetts, USA.
Partap U (1998). Foraging competition between Apis cerana and Apis mellifera and its impact on crop pollination. Asian bees and
beekeeping: progress of research and development. Proceeding of the Fourth Asian Apicultural Association International
Conference, Kathmandu, Nepal, 206–208.
Partap U, Shukla AN, Verma LR (2000). Comparative foraging behaviour of Apis cerana and Apis mellifera in pollinating peach and plum
flowers in Kathmandu Valley, Nepal. In Matsuka M, Verma LR, Wongsiri S, Shrestha KK, Partap U (Eds.) Asian Bees and
Beekeeping: Progress of Research and Development: proceedings of Fourth Asian Apicultural Association International
Conference, Kathmandu, March 23-28, 1998. 193–198. Published by Science Publishers, Enfield NH USA.
Radloff SE, Hepburn HR, Koeniger G (2003). Comparison of flight design of Asian honeybee drones. Apidologie 34: 353–358
Rao KS, Rao KL (2011). Management of floral resources to increase honey production through migration. In: Proceedings of State Level
Seminar on 'Awareness, Motivation & Technology Transfer for Development of Beekeeping in Andhra Pradesh,' January 8-9, 2011.
Published by the National Bee Board, Government of India.
Rinderer TE, Oldroyd BP, de Guzman LI, Wattanachaiyingchareon W, Wongsiri S (2002). Spatial distribution of the dwarf honey bees in
an agroecosystem in southeastern Thailand. Apidologie 33: 539–543.
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Ruttner F (1988). Biogeography and taxonomy of honeybees. Published by Springer Verlag, Heidelberg, Berlin.
Ruttner F, Maul V (1983). Experimental analysis of reproductive interspecies Apis mellifera L. and Apis cerana Fabricius. Apidologie 14:
309–327.
Sakagami SF (1959). Some Interspecific Relations Between Japanese and European Honeybees. Journal of Animal Ecology 28(1): 51–68.
Sharma HK, Gupta JK, Rana BS (2000). Diurnal activity of Apis cerana F. and A. mellifera L. on different flora during spring and honey
flow period. Pest Management and Economic Zoology 8(2): 151–154.
Verma LR (1993). Declining genetic diversity of Apis cerana in Hindu Kush-Himalayan Region. In: Connor LJ, Rinderer TE, Sylvester
HA, Wongsiri S. (Eds.) Asian apiculture 81–88, Published by Wicwas Press, Cheshire, Connecticut, USA.
Yang GH (2001). Chinese Honeybee. Published by China Agriculture Science and Technology Press, Beijing, China.
Yang G (2005). Harm of introducing the western honeybee Apis mellifera L. to the Chinese honeybee Apis cerana F. and its ecological
impact. Acta Entomologica Sinica 48(3): 401–406.
Yang MX (2009). Studies on Mixed–Species Colonies of Honeybees, Apis cerana and Apis mellifera. Ph.D. Thesis. Rhodes University,
South Africa.
Apis cerana Apis dorsata hive
Apis dorsata hive Apis dorsata
Apis florea hive Apis mellifera on Indian raspberry
Vol. No. 23(2) : March, 2017
10 ENVIS Newsletter
A. florea
A. andreniformes
A. dorsata
A. laboriosa
A. mellifera
A. koschevnikovi
A. nigrocincta
A. nuluensis
A. cerana
Small bees
Large bees
Medium sized bees
Phylogeny of the genus Apis (adapted from Oldroyd and Wongsiri 2006)
In the past ten years at least 12 alien pests have invaded Indian geographical boundaries or have been formally reported. The
spread of alien species is one of the greatest threats to the ecosystem. Classical biological control appears to be an important approach
in the management of these invasive species. International collaboration for the management of invasive species is of utmost
importance as exchange of resources can help in timely control of the pest.
The list of the invasive species in the last decade in the ascending order of their year of invasion/report is as follows:
Quadrastichus erythrinae Kim (Hymenoptera: Eulophidae)
In India, it was first recorded from Kerala (Faizal et al. 2006) and later from Karnataka and Tamil Nadu. The erythrina gall wasp forms
galls on the leaves, stems, petioles, and young shoots of Erythrina spp. As galls are formed on the newly emerging leaves, there is severe
reduction in the number and size of leaves besides complete cessation of growth. Affected trees appear to be scrawny with malformed and
crinkled shoots, suffer severe defoliation and ultimately die (Faizal et al. 2006).
Aprostocetus sp. (causalis-group), close to A. felix La Salle, Yang & Lin from Taiwan, is a potential parasitoid of erythrina gall wasp in
India. Species level confirmation requires extensive morphological studies including molecular analyses of all the Asian populations to
confirm the identity of the prevalent Indian populations.
Leptocybe invasa Fisher & La Salle (Hymenoptera: Eulophidae)
Leptocybe invasa is thought to originate from Australia. The wasp causes typical bump–shaped galls on the leaf midribs, petioles, and
stems of several Eucalyptus spp., particularly in young plantations and nurseries. Larvae form galls on the leaves and tender shoots of
Eucalyptus spp. Heavy galling prevents further development of the infested growth (Gupta & Poorani 2008 & 2009).
Quadrastichus mendeli Kim & La Salle, a solitary ectoparasitoid of L. invasa, known to be uniparental, develops on both young and
mature larvae of L. invasa. It was Introduced in India from Israel along with Selitrichodes kryceri Kim & La Salle, but the latter did not
establish.
Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae)
Since 2005, an introduced mealybug of the genus Phenacoccus has been causing serious damage to cotton (Gossypium hirsutum) in
north-western India. Hodgson et al. (2008) concluded from a comprehensive morphological study that there were no significant
differences in specimens from the Indian subcontinent compared to those from the Neotropics; and thus, considered the name P.
gossypiphilous to be a synonym of P. solenopsis.
ALIEN INVASIVE ARTHROPODS
Ankita Gupta*, Chandish R. Ballal and A. N. Shylesha
ICAR-National Bureau of Agricultural Insect Resources, Hebbal, Bengaluru-560024, Karnataka, India
*Corresponding author: ankitagupta.nbaii@gmail.com
