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Comparative Study of Apis cerena and Apis mellifera 

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Corresponding author:  Dharma Raj Katuwal  E-mail:
Honey bee 
Received: 13.05.2023 Received in revised form: 18.05.2023 
Accepted: 19.05.2023
This paper presents a comparative analysis of the two most important honeybees found worldwide, namely, Apis cerana (Asian honeybee) and  Apis mellifera (European honeybee). Both species are popular for their high-quality honey production and as effective pollinators. A. cerana is known for its hardiness, disease resistance, and ability to tolerate a wide range of temperatures. Additionally, A. cerana is smaller in size than A.  mellifera and exhibits a shyer nature. Both species have similar life cycles,  but A. cerana demonstrates greater hygiene awareness and is more capable of defending itself against potential threats or disturbances. This paper highlights the importance of understanding the unique characteristics of these two species, which can help inform management practices for beekeepers and aid in conservation efforts for these vital pollinators.


Honeybees (Hymenoptera: Apidae), which belong to  the genus Apis, are among the most well-known flying  insects found in terrestrial habitats (Koetz, 2013). There are nine currently recognized Apis honeybee  species worldwide, eight of which are native to Asia.  Apis mellifera is the only Apis honeybee species  outside of Asia (Koetz, 2013). Of the nine species of  Apis, only A. mellifera (the European or Western honeybee) and A. cerana (the Asian or Eastern  honeybee) have been domesticated for a long time,  and are of major commercial importance.

Most of  these species are limited within tropical and montane  zones in Southeast and South Asia, but the two species  have far broader ranges. A. cerana occurs as far north  as Japan and into the Middle East. European  honeybee A. mellifera, the most representative and  well-known honey bee, is native to Europe, Africa, and  most of Asia, but it has been introduced into the Americas and Oceania, where feral populations can be found (Breed, 2010). 

Due to their different ecological environment, they  have different behavioral and physiological  characteristics. Compared to A. mellifera, A. cerana has several distinguishing behavioral traits. It can  easily adapt to extreme weather conditions and has  long flight duration, effective grooming and hygienic  behaviors, and cooperative group-level defenses.

A  well-known behavior of A. cerana is aggregation when  a colony is exposed to dangers like predators or  intruders. In such situations, guard bees produce  alarm pheromones to communicate with other bees  about the danger. In addition, A. cerana provides  considerable economic benefits to the apicultural  industry through its high-quality bi-products, perhaps  more than A. mellifera (Park et al. 2015). A. mellifera has been subdivided into at least 20 recognized  subspecies or races (Koetz, 2013). Similar to A. mellifera, there are eight recognized subspecies of A.  cerana.
These subspecies tolerate a good range of  temperatures from cold, temperate, to tropical  ecosystems.  
Even though A. mellifera produces a large amount of  honey per colony, the indigenous A. cerana is  profitable over A. mellifera because A. mellifera requires intensive management, standardized  equipment, and larger foraging areas. A. cerana exhibits high tolerance to seasonal low temperatures  as compared to A. mellifera and as a result, it is the  first active honeybee on cool mornings in the northern  tropics (Corlett, 2010). 


A review of the published literature in national and  foreign journals, proceedings, reports, newsletters,  and books has been made in an effort to compare the  A. cerana and A. mellifera


Comparison of morphological characteristics of A.  melifera and A. cerana 

In general, A. cerana is smaller than A. mellifera. Apart  from a few conspicuous qualitative characteristics  which are mainly used to discriminate between the  very similar species A. cerana and A. mellifera, e.g.,  the radial vein of the hind wing, the tomentum on the  sixth tergite or the absence of chitinous plates on the  endo phallus, others are found which are also specific for A. cerana.

A. cerana have more prominent and consistent striping on their abdomen with even black bands across the abdomen, whereas A. mellifera tend  to have uneven black stripes with thinner stripes at  the front and thicker black stripes towards the rear  which makes it more yellow at the front and darker at  the back of the abdomen. The most reliable  morphological characteristic of A. cerana is the  extension of the radial vein on the hind wing, which is  absent in A. mellifera.

The differences between A.  cerana and A. melifera are most striking in the male  genitalia, while they are easily overlooked in the  female castes (Ruttner, 1988). The abdominal stripes  (tomenta) of A. cerana are more pronounced than  those of A. mellifera, and A. cerana workers have four  abdominal stripes, whereas A. mellifera workers have  three abdominal stripes.

Ecology and behavior

Foraging behavior 

Bees gather nectar and pollen from blossoms and  facilitate pollination at the same time. Nectar is later  turned into honey by the bees in a nest, to provide  energy in the form of carbohydrates for the colony.  Foraging is one of the distinctive behaviors of honey  bees which is the link between the honey bee colony  and the ambient environment. 

Table 1. Comparison of several specific characters  between A. cerana and A. melifera

Characters A. cerena A. melifera
Labrum – pigmentation All yellow or  brownAll dark or dark  with yellow mark
Tibia of hind leg – drone Grove (longitudinal)Round
Cubital index  (mean value) 4.40 2.30
Hooks on hind  wing (mean  value) 18.28 21.30
Endophallus  Chitinous  plates  Upper cornua   Fimbirate lobe Absent Three pairs Rosette-likePresent Rudimentary Feather-like

Source: (Ruttner, 1988)

On a single foraging trip, A. cerana foragers tend to  collect either pollen or nectar (not both) from a single  species of plant, continuing to collect pollen or nectar  from that plant throughout the day (Corlett, 2010).  Foraging ranges of A. cerana vary between different  studies, but maximum foraging ranges of 1,500 m to  2,500 m have generally been observed. In comparison,  A. mellifera tends to forage across much larger  distances, with maximum distances of over 10 km  (Beekman and Ratnieks, 2000).

Honeybees start and  finish foraging often depending on ambient  temperature, humidity and/or light levels of the day,  as well as the availability of floral resources. In  general, A. cerana tend to start foraging earlier than  A. mellifera, as it requires slightly lower temperatures,  light intensity and solar radiation levels to commence flight activity than A. mellifera. However, A. mellifera start foraging later than A. cerana because of its larger body size that requires a higher thoracic temperature,  and A. cerana are thus more adaptable to extreme  fluctuations in ambient temperature and long periods  of rainfall (Tan et al. 2012). A. cerana is more  industrious in collecting nectar from scattered  flowers, while A. mellifera worker has a stronger  foraging capacity of large flower patches (Feng et al.  2014). 

Fanning behavior

Honeybee colonies are able to maintain brood nest  temperature within the range of 33–36˚C, even when  the ambient temperature ranges from well below  freezing to above 45˚C (Fahrenholz et al. 1989).  Fanning bees stand on the landing platform at the  doorway to their hive and produce a current of air by  beating their wings, which serves as air-condition to  the hive. Since they remain for up to five minutes in  the same spot by gripping the floor with their claws,  their regular wing movements present a state of  ‘flight’ suitable for studying without fixing the animal  (Junge, 2006). 

One of the recognizable differences between A.  cerana and A. mellifera is the fanning position of  workers at the hive entrance in which A. cerana workers ventilate the hive by fanning away from the  entrance with their head, whereas A. mellifera fan  with their heads turning facing the entrance. The  entrances of hived bees are generally at the bottom of  the nest/hive. So, A. cerana workers face upwards,  whereas A. mellifera workers face downwards  (Ruttner, 1988).
A. cerana build multiple comb nests in dark cavities, although open nests (e.g., built underneath building  eaves) have also been observed. Combs are built  parallel with a uniform distance between the bee  spaces. Honey is stored in the upper and outer combs  adjacent to the cavity walls and the remaining comb  space is taken up by brood of various ages.

The number of combs in A. cerana nests varied from three  to fourteen combs. A. cerana cells are of two sizes:  generally smaller worker cells and larger drone cells  (Phiancharoen, et al. 2010). In comparison, the A.  mellifera worker cell is larger than that of A. cerana. A. cerana drone cells have a raised cap with a  distinctive pore at their apex. The size difference  between worker cells and drone cells is less noticeable  in A. cerana than in A. mellifera. Large conical queen  cells are built on the lower edge of the combs.  However, just like body size, worker cell size also  varies geographically and is larger in colder regions. 
The wax of A. cerana has a melting point of 65°C which  is about 2°C higher than that of A. mellifera (Ruttner,  1988). The drone cell is capped by the worker bees  with a wax cover and the drone larva spins the cocoon.  However, 1-3 days after completion of the cover, bees  start to remove the wax and then a yellowish, hard,  silky plate appears. Wherever the behavior of A.  cerana has been studied, no use of propolis was found  and the cracks in the hive are not sealed. However, a  brittle and light greyish mass is present on hive walls  and frames, which is not pure wax. 

Colony Defense

A. cerana is generally docile, gentle, mild, tolerant and  timid in defense behavior. A. cerana shows a number  of behavioral changes which prove to be very effective  against traditional enemies. Finally, A. cerana has  several unique responses to disturbances including  fast and sudden lateral body shaking of workers, the  production of a hissing sound, and heat balling.  

Hissing sound

When the beehive is disturbed or a certain enemy  attacks, it induces a sharp hissing sound that lasts  about 0.5 s. The sound is produced by a collective  quick closing of the wings over the body and the  abdomen jerks upward. The reaction is transmitted to  bees, besides immediate contact with the stimulating  agent, by body contact; it migrates over the comb or  the bee cluster with a velocity of 3 cm per second  (Koeniger and Fuchs, 1973). 

Group defense

Similarly, if it is attacked by powerful enemies such as  hornets, A. cerana bees do not counter-attack, as  most races of A. mellifera do. But, A. cerana forms  groups of 30 with the tip of the abdomen raised near  the entrance. The shy A. cerana stops flight activities  on a hornet’s arrival. The hissing sound is repeatedly  emitted in the hive. After this rapid retreat with no  solitary counter-attack the hornet usually relinquishes its attempt and leaves. If hornets persist in the attack,  they do not dare to capture bees out of the group. But,  if the hornet approaches too closely, it is seized  simultaneously at the legs and wings by several bees  and drowned in the mass of bees and killed. In most  of the observations, virtually no bees died during the  defense. The hornet attacks are stopped at the very  beginning.  
Heat balling is a unique defense of A. cerana to kill predatory hornets like Vespa simillima xanthoptera.  Several hundred bees surround the hornet in a tight ball and vibrate their thoracic muscles to produce heat. The A. cerana workers are able to raise the temperature inside the ball to an average of 46°C for approximately 20 minutes.

This temperature is high  enough to kill the hornet inside, but not high enough to kill the bees, who can tolerate temperatures up to  48 and 50°C. A. mellifera workers also will surround a  hornet, but they are not able to raise the temperature  as high as can A. cerana. Instead, A. mellifera workers  primarily sting the hornet and are less effective at  eliminating the hornet. So, the attacks on A. mellifera colonies by Vespa develop quite differently as it waits  for the attacking bees, seizes and kills them with its  strong mandibles and transports the corpses back to  the nest.

When the number of hunting wasps  increases with the decrease in the defense reactions  of the bee colony, the wasps kill one bee after the  other without carrying the corpses to their nest, until  the resistance ceases and they succeed in occupying  the beehive and eating or transporting pupae and  larvae. At the entrance of the occupied hive, the  territorial defense behavior of the wasps is observed  (Matsuura and Sakagami, 1973). A. cerana is the only  potential prey of V. mandarinia which has developed  an effective defense tactic. Group defense behavior  evidently represents a higher level of cooperation  than individual counter-attacks. 

Stinging behavior

A. cerana is in general less prone to sting than A.  mellifera. A. cerana has a reduced frequency of sting  apparatus autotomy. This is due to a change in  behavior by revolving movements after stinging  instead of straight runs and by strong muscles  anchoring the sting apparatus to the spiracle plates. A.  cerana uses its mandibles instead of the sting when attacking an enemy which resulted in the reduction of  the sting-protracting muscles.

Of all Apis species, A.  cerana has the least developed barbs on the lancets of  the sting (Koeniger et al. 1979). The venom of A.  cerana is identical to that of A. mellifera in the amino  acid sequence of the melittin, its main component.  Isopentyl acetate, an alarm substance was found in  worker bees in much lower quantities than in mellifera. 

Robbing and direct fighting 

When species come to overlap geographically and  compete for the same limited resources, either a  competitive exclusion or niche partitioning will occur.  It is possible that the ecological and behavioral  differences between A. mellifera and A. cerana will  result in sufficient niche partitioning so that both  species can co-occur successfully (Sharma, 2000).  Both species can also coexist if resources are not  limited. Floral resources and nest cavities are the two  most important resources for cavity-nesting  honeybees.

Competition for pollen and nectar may  occur on flowers, or they can attempt to rob honey  from other nests that can be of the same a different  species. Robbing bees enter another colony’s nest, kill bees  and take their honey store. The smaller the colony the  more susceptible it is to robbing (Partap, 2011).  Robbing occurs when floral resources are low, nectar  flow is interrupted, or a colony is weak or diseased. 

Interestingly, A. mellifera showed a much stronger  defense response than any of the Asian honeybees  (Breed et al. 2007). Studies on robbing behavior  between managed hives of two species kept at the same apiary showed that A. mellifera usually won by  killing A. cerana colony and taking over the foraging area although A. cerana initiated robbing during lean  times. No effective defense reactions are developed in  A. cerana to robbing attempts by A. mellifera.

There are no guard bees at the entrance so, intruders can pass easily without being inspected and also  sometimes robbed A. cerana bees were observed to feed the robber bees (Ruttner, 1988).

During the hot season A. cerana colony attacked by A. mellifera bees  usually loses all its stores and then absconds or dies.  Overall, evidence suggests that A. cerana is a weak  nest defender and competitor as compared to A.  mellifera. Compared with A. cerana, A. mellifera has a larger body size, longer flight range and stronger  defensiveness. A. mellifera thus shows advantages in  nectar robbing, nuptial flights and mating and disease transmitting (Tan et al. 2012). 

Brood development

The development of A. cerana is similar to that of Apis  species in general, and that of A. mellifera in  
particular. A. cerana brood development is slightly  faster than that of A. mellifera, except for A. cerana queens (Table 2) (Koeniger et al. 2010). However, it  seems doubtful that this slight difference will affect  competition or invasiveness. Like in other cavity nesting species, the larva’s brood cell is capped by the  worker bees just before the last of the five larval  instars. 

Table 2. Duration of the life cycle (days) of different castes of A. cerana and A. mellifera 

Stage Worker Drone Queen
A. cerana A. mellifera A. cerana A. mellifera A. cerana A. mellifera
Egg to larva 3
Larva to pupa 4-5.5 5
Pupa to adult 11 12 14 14 6-7.5 5
Total 19 21 23 24 13-16 13

Source: Koeniger et al. 2010 

Swarming and absconding

Reproductive swarming and absconding are the two  types of swarming in honeybees. Reproductive  swarming involves the splitting of a colony and  movement of the old queen with more than 70 % of  the colony to a new nest site, while the new queen  stays with the remaining colony and all its resources in  the old nest site. It generally occurs when conditions  are favorable and floral resources are abundant  (Chinh et al. 2005). There are two types of absconding

i. Seasonal absconding/migration: movement of the  whole colony due to resource depletion, declining  nest site quality 
ii.Disturbance-absconding: acute disturbance through  natural causes like fire, flooding, or anthropogenic like  intervention by beekeepers.

Migration is resource-related and is a seasonal movement of tropical honeybee colonies without any reproduction. It maximizes the colonization of new areas and provides a spatial refueling cycle clearly driven by reselection. To migrate/abscond, bees must have sufficient flight fuel and energy reserves so that,  they can construct new combs at a new site. 

Absconding colonies typically expand honey,  accelerate wax production, reduce oviposition and  consume eggs and young larvae so conserving protein.  Absconding/migration may be beneficial to the  survival, dispersal and propagation of honeybees, but  imposes serious difficulties for beekeeping in the  tropics (Hepburn and Radloff, 2011) Honeybees prepare for the move (lasting days to weeks) prior to  moving, when foraging, reduce honey and brood  levels during seasonal absconding but no such  preparation occurs before disturbance absconding.

In general, tropical honeybees including African strains  of A. mellifera, are more prone to absconding than  temperate species due to the change in temperature,  humidity, and resource levels. This means tropical  honeybees are able to move throughout the year in  response to change or disturbance, and to follow the  honey flow, both of which increase fitness and survival  (Ruttner, 1988). 

Seasonal absconding 

Seasonal absconding is directly related to resource  depletion and adverse environmental conditions of  the current location. A. cerana don’t store large  amounts of honey so they have no sufficient stores to  last for long. So, they move continuously to find better  conditions elsewhere during periods of high temperatures and dry seasons, after the abatement of  prolonged heavy rains (Hepburn and Radloff, 2011). 

Absconding has also been found highest in areas with  high environmental uncertainty like drought, and  when nest cavities are too small for the growing  colony. However, studies on A. cerana have also  observed absconding regardless of colony size,  congestion, or food supply (Hepburn, 2011) or without  an apparent external cause.

A. cerana prepare for migration by decreasing the  numbers of pollen-carrying workers, reducing brood feeding and rearing, and reduced predator and  parasite defense, decreasing honey and pollen stores,  eggs which leads to large changes in colony  demography (Pokhrel et al. 2006). A. cerana abscond  less often than open-nesting Asian honeybee species  but more often than temperate A. mellifera.  Temperate A. mellifera, especially wild colonies, may  abscond for to the same reasons as tropical  honeybees due to depleting resources and starvation,  predation, disturbance, adverse environmental  conditions, and disease/parasitism (Ruttner, 1988). 

Predation (Disturbance absconding) 

Tropical honeybee species are under more severe  predation pressure than temperate honeybees.  Predation is an important and powerful force in the  evolution of Asian honeybees, shaping choice of nest  site, nest architecture, population size, worker  morphology, and behavior.

Natural predators of A.  cerana include wasps, ants, vertebrates (tiger, human,  birds etc.), and hornets, which tend to prey on  foragers but also at times attack colonies. In cavity nesting Apis species, the main defense against  predators is living in a protected cavity with a small  entrance that can be easily guarded. Colony defense  behaviors include abdomen shaking, hissing (through  wing vibrations), group defense (including grasping,  pulling, and biting, killing by overheating and/or  asphyxiation), and stinging (Ruttner, 1988). 

Reproductive swarming in A. mellifera occurs when  floral resources are abundant and a colony is  performing well (Chinh et al. 2005). Soon after a  swarm has left the old nest, A. mellifera settles tens of  meters away and scouts will start searching for  suitable nest sites. Similarly, A. cerana also settles 20  ± 30 m away from the old nest for several days and  then moves to the new nest site. A. mellifera colonies  are prevented from swarming by good colony  management, removing new queen cells, re-queening  and using queen excludes.

Wild, temperate A.  mellifera, however, swarm nearly every year and sometimes up to three times per seasonal cycle when  resources are highest. Swarming of A. cerana is highly variable and depends on the geographic location and climate. A. cerana can swarm several times a year  (Ruttner, 1988). According to Koeniger et al. (2010),  swarming will start when a colony reaches 20,000 bees, with an average of eight swarms per colony.

The timing of swarming has been found to vary from no  seasonal rhythm, biphasic, to distinct times of the year  (Hepburn, 2011).When foraging conditions are good over a long time,  swarming will occur more frequently resulting in the  asynchronous production of queens and drones  (Chinh et al. 2005). When foraging conditions are good  only at certain times of the year (e.g., spring and summer in temperate zones), swarming will occur  during those specific times, and swarming and the  production of queens and drones will be synchronous  as seen in temperate A. mellifera (Hepburn, 2011). 

Diseases and Hygiene 
Where honeybee species coexist, they are bound to  interact in some way like robbing due to which  parasites and pathogens can be transmitted between species. Diseases and parasites have been introduced  from A. mellifera to A. cerana like the tracheal mites as well as Israeli acute paralysis virus and Varroa destructor from A. cerana to A. mellifera. A. cerana diseases include bacterial infections such as American and European foulbrood, protozoan and fungal  infections like Nosema ceranae and N. apis and  chalkbrood, and virus infections like Apis Iridescent  virus, Deformed wing virus, Thai sacbrood virus, black  queen cell virus etc. A. cerana parasites include Varroa  (V. destructor, V. jacobsoni, and V. underwoodi) and  tracheal mites (Acarapis woodi), as well as non parasitic mites (Kojima et al. 2011). Varroa jacobsoni Oudemans is widespread throughout the cerana area. 

However, no severe damage is caused to A. mellifera due to absence of a particular host-parasite relation  (Ruttner, 1988). A. cerana workers were found to  clean themselves more thoroughly than A. mellifera.  In addition, infected brood is either removed before capping (e.g., larvae infected with American foulbrood  or worker brood with Varroa), or is entombed (e.g.,  drone larvae infected with Varroa) (Rath, 1999). 

Experiments showed that immediate cleaning  behavior in A. cerana is due to the presence of Varroa  semiochemical compounds. A. cerana are regarded as  hardy and disease-resistant than A. mellifera, making  it a better species in many poorer areas of Asia as A.  cerana requires less management and treatment for  diseases (Russo et al. 2020).


Hence, Apis cerana, an Asian honeybee and Apis  mellifera, European honeybee are the two most  important bee hives found in the wide range of the  world. Both the species are cavity-nesting bee hives  and are popular for their high-quality honey  production and a good pollinator. A. cerana can  tolerate a wide range of temperatures and are of shy  nature, hardy and disease resistant as compared to A.  mellifera. A. cerana is smaller than A. mellifera in  average and have a similar life cycle. A. cerana are  capable of defending themselves against enemies or  disturbance and are conscious about their hygiene in  comparison to A. mellifera.



We would like to express our sincere gratitude to the  reviewers for their insightful comments and  suggestions, which significantly contributed to the  improvement of this manuscript. Our thanks are also  due to all the researchers and authors whose works  have been referenced and cited in this review, as their  contributions have greatly enriched the content and  depth of this article. 


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