Apis cerana

Eastern honey bee
Asiatic honey bee - Khao Yai National Park, Thailand
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Suborder: Apocrita
Family: Apidae
Subfamily: Apinae
Genus: Apis
Subgenus: (Apis)
Species: A. cerana
Binomial name
Apis cerana
Fabricius, 1793
Map showing the range of Apis cerana
Range of Apis cerana

Apis cerana, or the Asiatic honey bee (or the eastern honey bee), is a species of honey bee found in southern and southeastern Asia, including China, Pakistan, India, Korea, Japan, Malaysia, Nepal, Bangladesh, Papua New Guinea and Solomon Islands. This species is the sister species of Apis koschevnikovi, and both are in the same subgenus as the western (European) honey bee, Apis mellifera.[1][2][3][4] A. cerana is known to live sympatrically as well with Apis koschevnikovi within the same geographic location.[5] Apis cerana colonies are known for building nests consisting of multiple combs in cavities containing a small entrance, presumably for defense against invasion by individuals of another nest.[6] The diet of this honey bee species consists mostly of pollen and nectar, or honey.[7] Moreover, Apis cerana is known for its highly social behavior, reflective of its classification as a type of honey bee.[4]

Taxonomy and Phylogeny

Danish zoologist Johan Christian Fabricius described Apis cerana, also known as the Eastern or Asian honey bee, in 1793.[2] The genus name Apis is Latin for “bee.” The Asiatic honey bee is of the Apidae family, one of the most diverse families of bees, including common honey bees, carpenter bees, orchid bees, bumblebees, cuckoo bees, and even stingless bees.[8]

In the past, there has been discussion that Apis cerana and Apis mellifera are simply distinct races of the same species. This is essentially due to overwhelming similarities in both morphology and behavior, as both are medium sized bees (10-11mm) that generally build multiple comb nests inside cavities. Other honey bee species, including the giant honey bees Apis dorsata and Apis laboriosa, generally construct nests consisting of a single comb in open areas.[9] However, despite the striking similarities between Apis cerana and Apis mellifera, there is evidence to suggest that these two species are quite distinct; for example, mating between these species does not produce offspring. In addition, while Apis mellifera colonies can reach sizes of up to 50,000 or more individuals, Apis cerana colonies are relatively small, with only around 6,000 to 7,000 workers.[9] Morevoer, Apis cerana is found predominantly in the Eastern Asian region of the world, while Apis mellifera is found predominantly in the Western European and African region of the world.[9] For these reasons, it has now been concluded that these are in fact two separate species, contrary to prior beliefs.

Subspecies

(following Engel, 1999).

Eight subspecies of A. cerana are currently recognized. Of these, two subspecies are predominant and used for apiculture in India: A. c. cerana and A. c. indica. These species are similar to Apis mellifera except in color. A. c. indica has black stripes on its abdomen and lives close to hilly areas and is sometimes seen in plains regions. A. c. cerana has yellow stripes on its abdomen and is habituated to plains regions of India. Apis mellifera can be readily distinguished from A. cerana due to its slightly larger size.

Description and Identification

The physical characteristics of individuals of the Apis cerana species are very similar to those of other species in the Apis genus. The individuals in this genus are defined by long, erect hairs that cover the compound eyes and assist in pollen collection, strongly convex scutellum, and a jugal lobe in the hindwing.[4] Adult Apis cerana are black in color, with four yellow abdominal stripes.[4] There are also distinctions between worker bees, queens, and drones. Worker bees are characterized by a pollen press on the hind leg to transport pollen, as well as a stinger in the place of an organ for laying eggs.[4] Queens, which are the reproductive females, are typically larger than worker bees due to their enlarged reproductive organs.[4] Drones, which are the males of the species, are defined by larger eyes, lack of a stinger, and a blunter abdominal shape.[4]

Distribution and Habitat

Apis cerana encompass a wide range of climatic zones including moist tropical rainforests, wet-dry tropical savannas, mid-latitude steppes, dry mid-latitude grasslands, moist continental deciduous forests, and taigas.[10] The species naturally occurs in an East Asian landscape, though predominantly found in China, the Philippines, New Guinea, the Soloman Islands, and recently even near Australia.[10] Although the species was naturally clustered in East Asia, it has now expanded to various regions across the world as a result of human interference, with particular concern about its spread into Australia as nests are found in a variety of environments, including both natural and man-made.[10]

Nest

As a type of honey bee, Apis cerana must collect and store around a third of its nectar in a concentrated form in order to ensure an adequate supply for consumption during the harsh winter.[11] The nest is multi-combed and somewhat insulated, allowing the species to achieve optimal nest temperatures and thereby optimize foraging at early hours. The nest itself is a simple vertical comb made from wax secreted by workers, while the thermoregulation is generated by fanning and water evaporation from water collected in the field.[4] This large-scale storage tactic requires the construction of a stable and sturdy comb structure for such honey storage. However, as the production of beeswax for a comb is an energetically expensive endeavor, colonies do not generally build the entire structure very early on.[11] Instead, the colony builds a smaller set of combs that satisfies the initial storage needs. Upon moving into the new nest, the colony will then continue to enlarge the combs until there are enough storage units to sufficiently account for all of the required honey.[11] In general, the initial set of combs may account for around 20,000 cells, while the final, completed structure may contain up to 100,000.[11] These nests are typically large enough to host around 6,000 to 7,000 individuals and are found in a wide range of external environments.[9]

Nest thermoregulation

A. cerana maintains internal hive temperatures with a precision similar to that of A. mellifera, using similar mechanisms. A. cerana maintain body temperatures in a range of 33–35.5 °C even while ambient temperatures vary between 12 and 36 °C. This mechanism clearly shows them to possess effective nest thermoregulation systems. During summer, A. cerana employs evaporative cooling, where the worker bees cluster outside the nest in hot weather and fan their wings, thus removing excess heat and moisture from the nest and decreasing the hive temperature.

Behavior

Colony Cycle

The colony of Apis cerana, a typical honey bee, consists of several thousand female worker bees, one queen bee, and several hundred male drone bees. The colony is constructed inside beeswax combs inside a tree cavity, with a special peanut-shaped structure on the margins of the combs where the queens are reared.[11]

The colony’s annual cycle in cold temperature regions begins shortly after the winter solstice, when the colony raises the core temperature of its cluster to about 34 degrees Celsius and starts to rear brood. At first, only around 100 bees are produced, but several thousand bees are developing by early spring. By late spring, the colony will have already attained full size, and will begin to reproduce. The colony then rears several new queens, and divides itself with about half the workers plus the old queen once the new queens have nearly matured. This new swarm then flies to a new tree branch, explores nest cavities, and then directs the other bees to the new site once satisfied with the location.[11]

During the remainder of summer and into the fall, the colonies in the new locations build combs, rear brood, and gather food to quickly rebuild their populations and food reserves prior to the arrival of winter.[11]

Division of Labor

As a social species, Apis cerana colonies contain divisions of labor depending on what each member of the group is specialized to perform. There is generally only one queen bee whose sole responsibility it is to lay eggs; therefore, she is the mother of all the workers present in the colony. Apart from the queen bee, the remaining female bees are known as the worker bees, as these individuals perform all the tasks necessarily to maintain the hive including tending to the eggs, larvae, and pupae, foraging for food and water, cleaning the facility, and producing honey. These tasks are divided among the female worker bees by a factor of age. The remaining individuals are the males, known as the “drones,” whose only responsibility is to mate with queen from another colony; therefore, drones are solely produced during the reproductive season.

Communication

The principle method of communication is the waggle dance, performed primarily when a worker bee discovers a rich source of pollen or nectar and wishes to share this knowledge with her fellow nest-mates. The waggle dance occurs deep inside the colony’s hive, where the worker bee performs a brief reenactment of the recent journey to a patch of flowers. Neighboring bees observe and learn this dance and can then follow the same pattern, utilizing the odor of the flowers to fly in a certain path and arrive at the same destination. The bees following the informed worker bee will extend their antennae towards the dancer in order to detect the dance sounds, as the frequency of the bee’s antennae closely matches the vibration frequency of its wings.[11] The overall direction and duration of each waggle is closely correlated with the direction and distance from the flower patch being described.[11]

Mating Behavior

Within the honey bee colony, a queen bee typically mates with 10 or more males.[11] This extensive mating is performed in an effort to secure a great range of genetic variation in her colony to cope with diseases, as well as respond to nectar sources and a wide range of external stimuli.[11] Apart from the queen bee, the only other sexual members of the society are the male drones, whose only function is to simply mate with the queen, after which they will die.[9]

The exact time and place of Apis cerana mating is specific to the subspecies, often varying by local environment. For instance, in Sri Lanka, Apis cerana males typically aggregate beside a tree canopy as opposed to above a tree as is found in the Apis cerana subspecies of Japan. The most significant factor in determining mating time, however, is not ecological conditions, but rather the presence of drones of other species. Mating time decreases as the number of external species present increases.[4]

Reproductive swarming

In A. cerana, reproductive swarming is similar to A. mellifera. A. cerana reproductive swarms settle 20–30 m away from the natal nest (the mother or primary colony) and stay for a few days before departing for a new nest site after getting information from scout bees.[11] Scout bees search for suitable cavities in which to construct the swarm’s home.[11] Successful scouts come back and report the location of suitable nesting sites to the other bees by performing communication dances on the surface of the swarm cluster in the same way as for food sources.[11]

Absconding behavior

Apart from reproductive swarming, A. cerana has migration and absconding behavior, abandoning the current nest and building a new nest in a new location where an abundant supply of nectar and pollen is available. These bees usually do not store great amounts of honey, so they are more vulnerable to starvation if a prolonged shortage of nectar and pollen occurs. Absconding will start when not enough pollen and nectar are available. After the last brood emerges, the adult bees fill their honey stomachs from the hive's stores and swarm to establish a new nest at a new location. A. cerana has more absconding behavior than A. mellifera.

Life History

The development of worker bees in a colony is typical of that for any insect that undergoes complete metamorphosis as it includes the four stages of egg, larva, pupa, and adult. The embryo grows inside the egg for 3 days, consuming the protein-rich egg yolk.[11] Then it undergoes an 8-day larval stage, which is an intense feeding state involving honey, pollen, and brood food supplied by the adult bees.[11] Finally, there is construction of a wax pupa which then matures and gnaws through the wax cap of the cell to emerge as a young bee.[11]

Kin Selection

Genetic Relatedness Within Colonies

As one queen generally mates with over a dozen males, the genetic relatedness of the colony is biased and represents haplodiploid sex determination. If the queen bee lays unfertilized eggs with no paternal genetic contribution, the eggs will develop into drones. If the queen bee lays fertilized eggs with both maternal and paternal genetic contribution, the eggs will develop into females. In this system, virgin queens sharing the same father will have a genetic relatedness of .75 and those of different fathers will have a genetic relatedness of only .25.[11] The females workers in the colony are related to the queen’s sons by a genetic relatedness of .25.[11] Such biasing results in the genes of some female worker bees being represented disproportionately in the virgin queens.[11]

Worker-Queen Conflict

Conflict may arise between workers and bees as female workers attempt to increase the propagation of their genes by biasing their queen-rearing efforts in favor of virgin queens sharing the same father. Although female worker bees do possess ovaries and can essentially produce viable eggs, this potential is almost never realized as long as the colony is ruled by a dominant queen. Therefore, the probability of personal reproduction by a worker bee is exceedingly low. “Worker policing,” which is the mutual prevention of reproduction by workers, could be the reason behind the conscious non-reproduction of female worker bees.[11] In other words, their fertility is controlled by queen signals. The queen honeybee informs workers of her presence by pheromones that she secretes from her mandibular glands. These signals are acquired by workers in close vicinity of the queen and then spread to other workers in the colony, mainly by body contacts. In presence of queen pheromone signals, the vast majority of workers refrain from activating their ovaries. Therefore, due to factors of genetic relatedness, Apis cerana often try to prevent other workers in her colony from reproducing, either by destroying worker-laid eggs, or by showing aggression towards workers attempting to lay eggs through worker policing.[11]

Interaction with Other Species

Diet

Adult worker bees predominantly feed on pollen and nectar or honey, though the nutritive value of pollen varies depending on the plant. Mixed pollens possess a high nutritive value and actually supply all the necessary materials for proper development of young animals. However, when dried, pollen quickly loses its nutritive value.[7]

In addition to feeding themselves, bees also feed each other through a process known as “food transmission.” Moreover, workers may also obtain food from the queen, while drones acquire food by ingesting material regurgitated by other drones. Queens themselves are fed larval food by the workers during their wintering season, thereby neither feeding on nor being fed honey.[7]

Water Requirement

In addition to food requirements for diet, water also plays a key role in the growth and development of adult bees. In fact, the form of food has little to no influence on the longevity and life expectancy of the bee as long as there is still ready availability of water.[7] This idea was further supported through experimental means concerning queen bees isolated in separate cages. Both groups were fed sugar candy as a control measure; however, the group that was fed water in addition to the sugar candy lived an average of two weeks or more compared to the other group which only survived a matter of three to four days.[7] Moreover, the importance of water intake for an adult bee’s survival can be further understood through examination of diet and behavior during prime seasons, during which colonies of bees consume large amounts of water in order to dilute honey produced as well as to regulate temperature in the nest.[7]

Predators

Vespine wasps, endemic to Southeast Asia, are a major predator for Apis cerana, predominantly at their colonies throughout Southern Asia. This hawking predation is especially fierce during the autumn season when the wasps are most populous, predominantly during the morning and afternoon. This method involves the wasps taking up a position in front of the beehive, while facing outwards away from the entrance towards returning foragers.[13] Vespine wasps attempt to attack the honey bee quarry in an effort to gain provisions to aid in the development of their own offspring.[13]

Defense

As the Vespine wasps approach the entrance to the honeybee nest, more guard bees are alerted, which in turn increases their probability of being killed by heat-balling bees. Heat balling is a unique defense system in which several hundred bees surround the wasp in a tight ball and vibrate their muscles in an effort to produce heat and effectively kill the wasp inside.[13] Alternatively, however, in the presence of a wasp, the bees may also just withdraw into their nests and await the heat-balling circumstances to develop naturally. Furthermore, other bees may just decide to fly away as an evasive measure in times of conflict, often altering their specific flight styles in order to avoid predation.[13]

Thermal defense

When an A. cerana hive is invaded by the Japanese giant hornet (Vespa mandarinia), about 500 Japanese honey bees (A. cerana japonica) surround the hornet and vibrate their flight muscles until the temperature is raised to 47 °C (117 °F), heating the hornet to death, but keeping the temperature still under their own lethal limit (48–50 °C).[12]

Wing Shimmering

Although both Apis mellifera and Apis cerana suffer from predation from Vespine wasps, one defense mechanism unique to Apis cerana is “wing shimmering”.[13] During this period of self-defense, bees collectively execute carefully timed waves of shimmering of their wings when approached by predators such as Vespine wasps.[13] This appears to serve as a distraction technique of visual pattern disruption that results in confusing predators.[13] As a result, predators are unable to continue attacking additional bees, and are therefore unsuccessful in this endeavor.

Pests

Apis cerana is the natural host to the mite Varroa jacobsoni and the parasite Nosema ceranae, both serious pests of the western honey bee.[14] When first discovered near Beijing, China, in 1994, it was originally thought that Nosema ceranae was restricted to Apis cerana within in the East Asian region.[15] However, it has now been confirmed that this parasite species is actually present in colonies of Apis mellifera as well, both in Taiwan as well as Spain, though the origins of its arrival in Europe are still unknown.[15] Bees infected with Nosema ceranae suffer reduced lifespans as well as increased mortality in the winter and poor buildup and reduced honey yield in the spring.[15] Having coevolved with these parasites, A. cerana exhibits more careful grooming than A. mellifera, thus has an effective defense mechanism against Varroa that keeps the mite from devastating colonies. Other than defensive behaviors such as these, much of their behavior and biology (at least in the wild) is very similar to that of A. mellifera.

Genetic database

The Biomodeling Laboratory at Seoul National University has constructed an Asian honey bee transcriptome database using a next-generation sequencing technique (Illumina hiseq2000 and GS-FLX 454 technology). This database interface will support researchers to get the molecular and sequence information about A. cerana more easily.view

Gallery

References

  1. ↑ BIODIVERSITY OF HONEYBEES, M.R.Srinivasan, Department of Agricultural Entomology - Tamil Nadu Agricultural University accessed Jul 2010
  2. 1 2 Engel, M.S. (1999) The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: Apis). Journal of Hymenoptera Research 8: pp. 165–196.
  3. ↑ Photos of Apis cerana
  4. 1 2 3 4 5 6 7 8 9 Oldroyd, Benjamin P.; Wongsiri, Siriwat (2006). Asian Honey Bees (Biology, Conservation, and Human Interactions). Cambridge, Massachusetts and London, England: Harvard University Press. ISBN 0674021940.
  5. ↑ Koeniger, N.; Koeniger, G.; Tingek, S.; Kelitu, A. "Interspecific rearing and acceptance of queens between Apis cerana Fabricius, 1793 and Apis koschevnikovi Buttel-Reepen, 1906". Apidologie 27 (5): 371–380. doi:10.1051/apido:19960505.
  6. ↑ Nanork, P., et al. "Social parasitism by workers in queenless and queenright Apis cerana colonies." Molecular ecology 16.5 (2007): 1107-1114.
  7. 1 2 3 4 5 6 Haydak, Maykola H. "Honey bee nutrition." Annual review of entomology 15.1 (1970): 143-156.
  8. ↑ Michener, Charles Duncan. The bees of the world. Vol. 1. JHU Press, 2000.
  9. 1 2 3 4 5 Winston, Mark L. The biology of the honey bee. Harvard University Press, 1991.
  10. 1 2 3 Radloff, Sarah E., et al. "Population structure and classification of Apis cerana." Apidologie 41.6 (2010): 589-601.
  11. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Seeley, Thomas D. The wisdom of the hive: the social physiology of honey bee colonies. Harvard University Press, 2009.
  12. 1 2 Apis cerana' "cooking" a hornet to death video
  13. 1 2 3 4 5 6 7 Tan, Ken, et al. "Bee-hawking by the wasp, Vespa velutina, on the honeybees Apis cerana and A. mellifera." Naturwissenschaften 94.6 (2007): 469-472.
  14. ↑ Ritter, Wolfgang Nosema ceranae Albert Ludwigs University of Freiburg
  15. 1 2 3 Paxton, Robert J., et al. "Nosema ceranae has infected Apis mellifera in Europe since at least 1998 and may be more virulent than Nosema apis."Apidologie 38.6 (2007): 558-565.
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