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We review the diversity of honeybees in India and their documented interactions in this brief overview published in ENVIS Newsletter on Invasive Species in March 2017 (Vol. 23, Issue 2)
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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).
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).
1 1 2
Manjishtha Bhattacharyya *, Susanta Kumar Chakraborty and Sankar Kr. Acharya
1Department of Zoology, Vidyasagar University, Midnapore 721102
Email: *,
2Department of Agricultural Extension, Bidhan Chandra Krishi Viswavidyalay, Mohanpur, Nadia 741252.
*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).
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
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.
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.
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
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
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8ENVIS Newsletter
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.
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.
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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:
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B 169: 107–112.
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Research 11(1): 65–69.
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flowers in Kathmandu Valley, Nepal. In Matsuka M, Verma LR, Wongsiri S, Shrestha KK, Partap U (Eds.) Asian Bees and
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Apis ceranaApis 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
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.
Ankita Gupta*, Chandish R. Ballal and A. N. Shylesha
ICAR-National Bureau of Agricultural Insect Resources, Hebbal, Bengaluru-560024, Karnataka, India
*Corresponding author:
... Yet studies on the ecology of invasive species are better known in developed countries than developing one. In India, very scanty research studies have been undertaken on bioinvasion in general and aquatic invasive species in particular (Maheswari 1965;Maiti and Guhabakshi 1981;Ramkrishnan 1991;Bhakat et al. 2004;Paria et al. 2017;Bhattacharrya et al. 2017). Almost no such works are available from Indian coastal-mangrove ecosystem excepting scanty reports from Kerala coast of India (Ravinesh and Bijukumar 2016; Bijukumar and Raj 2017;Rajagopal et al. 1989) and of West Bengal coast (Bhakat et al. 2004). ...
... Researchers have estimated India's economic loss due to bio-invasion as to be 20% of its annual GDP, as opposed to less than 1% of GDP of the USA in the same year (Pimental et al. 2001). This is why invasive species are being increasingly regarded with animosity and alarm (Paria et al. 2017;Bhattacharrya et al. 2017). The change in land use and habitat alteration is accentuating species to invade areas which ultimately threaten local and specifically adapted faunal species (Perrings et al. 2010). ...
Full-text available
Impact of Bioinvasion on Coastal biodiversity of West Bengal coast, India
Full-text available
The sympatric congeners Apis dorsata, A. cerana, A. andreniformis and A. florea were observed foraging for pollen on the noctumally-dehiscent king palm (Archontophoenix alexandrea). The larger A. dorsata and A. cerana foraged earliest but in low numbers, presumably exploiting the resource at its most productive time. The smaller A. andreniformis and A. florea followed in large numbers. Although there was minimal separation of A. andreniformis and A. florea foragers in either space or time, no aggressive interactions between the species were observed.
The comparative aggressiveness of a yellow Californian strain of A. mellifera and of the hill variety of A. cerana indica was studied. It was found that, in general, A. mellifera was more aggressive than A. cerana. It is expected that under favourable natural conditions, A. mellifera will increase rapidly even in the presence of A. cerana indica.
Competition experiments at an artificial feeding dish revealed interactions among the species according to body size: Trigona attacked mainly A. florea and A. cerana; A. florea reacted to A. cerana, and A. cerana attacked A. dorsata. In exclusion experiments, smaller species proved more successful than larger ones.Altogether 51 comb samples with honey and pollen stores were examined microscopically; 46 pollen types recorded are listed, and most of them identified. The average number of different pollens in individual samples, and the absolute number in all samples from one bee species, were highest for T. iridipennis and lowest for A. dorsata. There was a correlation between the aggressiveness of a bee species and the versatility of the pollens in food stores. For some of the most common bee plants, correlations were found between body size of the bee species and the frequency of occurrence of the pollen grains in its food stores.Competition for food is not a limiting factor for the co-existence of the four bees studied. The disadvantage of the limited flight range of a small species is compensated by a more aggressive behaviour, which allows it to defend food sources in its smaller territory, and then to prevent disastrous competition.
Bee conservation is vital for the functioning of plant communities and human welfare. Unfortunately, bee population is declining in many parts of the world resulting in pollination crisis. The locally adapted strains, subspecies and ecotypes of Honeybees suffer less from elevated losses than non native bees Therefore, their conservation as genetic resource for breeding of disease and stress resistant strains is essential. Besides, a full understanding of origin and distribution of bees is very crucial for understanding how and when these adaptations arose. Understanding the evolutionary relationships of these bees would provide a basis for behavioural studies within an evolutionary framework, illuminating the origins of complex social behaviour, such as the employment of dance and sound to communicate the location of food or shelter. In addition to a global phylogeny, would also provide estimates of divergence times and ancestral biogeographic distributions of the major groups. In this chapter we discuss the origin, taxonomic composition and patterns of distribution of honeybees.
The excess power index (integrating body dry mass, thorax-to-body dry mass and wing surface area) was compared in drones of seven Asian Apis species. There are two statistically distinct groups of drones: drones of the dwarf honeybees form one class, all other Asian species belong to the second. Drones of dwarf honeybees have a 36% ergonomic advantage in power availability and 20% advantage in available excess power over all other drones. Comparisons of flight dimensions between conspecific workers and drones show a highly statistically significant sexual dimorphism for flight. Although drones of all seven tested species are always bigger than their workers their excess power index is some 15% better. It is suggested that prowess of flight in drones is driven by the need to compete and mate with queens flying high in the air while worker bees forage nectar and pollen on flowers.
1. Some observations on the interspecific relations between two species of honeybees inhabiting Japan, Apis mellifera L. (introduced) and A. cerana cerana Fabr. (endemic), are described. 2. In general, A. cerana is more tolerant and less aggressive than A. mellifera. 3. It is suggested that robbery by A. mellifera plays an important role in the decrease of A. cerana.