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Apis cerana

Apis cerana, the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of Apis koschevnikovi and both are in the same subgenus as the western (European) honey bee, Apis mellifera. A. cerana is known to live sympatrically along with Apis koschevnikovi within the same geographic location. 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. The diet of this honey bee species consists mostly of pollen and nectar, or honey. Moreover, Apis cerana is known for its highly social behavior, reflective of its classification as a type of honey bee.

Taxonomy and phylogeny
Danish zoologist Johan Christian Fabricius described Apis cerana, also known as the eastern or Asian honey bee, in 1793. 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. 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. • Apis cerana heimifeng Engel (black Chinese honey bee) - highlands in central China • Apis cerana indica Fabricius (Indian honey bee) - southern India, Sri Lanka, BangladeshApis cerana japonica Fabricius (Japanese honey bee) - JapanApis cerana javana Enderlein (Javan honey bee) - Java to East TimorApis cerana johni Skorikov (Sumatran honey bee) - SumatraApis cerana nuluensis Tingek, Koeniger and Koeniger (Bornean honey bee) - BorneoApis cerana skorikovi Engel (= himalaya) (Himalayan honey bee) - the central and eastern Himalayan mountains (Ruttner, 1987) Recent genetic analysis, however, has determined that some of the subspecies described may have been inadvertent misidentifications of very similar sympatric species, including Apis koschevnikovi of Borneo and Apis nigrocincta of the Philippines. Apis cerana nuluensis of Borneo is also now generally considered to be a separate species, as Apis nuluensis. Radloff et al. (2010) have instead chosen to subdivide Apis cerana into six main statistically defined populations based on morphotypes ("morphoclusters"), instead of infraspecific ranks, which they argue were invalidly established and not biologically meaningful. These morphoclusters are: • 'Northern cerana''' (Morphocluster I) - extends from northern Afghanistan and Pakistan to northwest India, southern Tibet, northern Myanmar, China and into the Korean peninsula, far eastern Russia and Japan. They can be further subdivided into six subclusters: an "Indus" group (Afghanistan, Pakistan, Kashmir); a "Himachali" group (Himachal Pradesh, India); an "Aba" group (larger bees in southern Gansu, central and northern Sichuan, northern China and Russia); a central and eastern China subcluster; a "southern" cerana'' subcluster in southern Yunnan, Guangdong, Guangxi and Hainan; and a "Japonica" group in Japan, North Korea and South Korea. :*Synonyms: A. skorikovi, A. c. abansis, A. c. abanensis, A. c. bijjieca, A. c. cathayca, A. c. cerana, A. c. fantsun, A. c. hainana, A. c. hainanensis, A. c. heimifeng, A. c. indica, A. c. japonica, A. c. javana, A. c. kweiyanga, A. c. maerkang, A. c. pekinga, A. c. peroni, A. c. skorikovi, A. c. shankianga and A. c. twolareca • 'Himalayan cerana''''' (Morphocluster II) - extends from northern India, Tibet and Nepal. Has two subclusters: the "Hills" group (northeast) and the "Ganges" group (southwest). :*Synonyms: A. c. indica • 'Indian Plains cerana (Morphocluster III) - extends from the plains of central and southern India and into Sri Lanka; also known as the Plains cerana '. :*Synonyms: A. c. indica • 'Indo-Chinese cerana''''' (Morphocluster IV) - extends from Myanmar, northern Thailand, Laos, Cambodia and southern Vietnam. :*Synonyms: A. c. indica and A. c. javana • 'Philippine cerana''''' (Morphocluster V) - restricted mostly to the Philippines, excluding Palawan. Has three subclusters, the "Luzon" bees, "Visayas" bees and "Mindanao" bees, with the latter two being more closer morphometrically than the "Luzon" bees. A population is also found in central Sulawesi. :*Synonyms: A. philippina, A. c. philippina and A. c. samarensis • 'Indo-Malayan cerana''''' (Morphocluster VI) - extends from southern Thailand, Malaysia, Indonesia and Palawan (the Philippines). Has three subclusters: one in Palawan, another in northern Borneo and Kalimantan and another in Java, Bali, Irian Jaya, Sulawesi and Sumatra. :*Synonyms: A. cerana, A. indica, A. javana, A. c. johni, A. lieftincki, A. peroni, A. vechti linda and A. v. vechti Latest 2023 list of sub species of APIs cerana 1) Apis cerana cerana 2) Apis cerana kashmirensis 3) Apis cerana japonica 4) Apis cerana hainana 5) Apis cerana abansis 6) Apis cerana guidensis 7) Apis cerana skorikovi 8) Apis cerana Taiwanesis == Description and identification==
Description and identification
The physical characteristics of Apis cerana individuals are very similar to those of other species in the genus Apis. 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. Adult Apis cerana are black in color, with four yellow abdominal stripes. 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. Queens, which are the reproductive females, are typically larger than worker bees due to their enlarged reproductive organs. Drones, which are the males of the species, are defined by larger eyes, lack of a stinger, and a blunter abdominal shape. ==Distribution and habitat==
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. 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 invasive potential in Australia as nests are found in a variety of environments, including both natural and man-made (see below). ==Behavior==
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. 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. 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. 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 beehive 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 a queen from another colony; therefore, drones are solely produced during the reproductive season. Communication The principal 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. The overall direction and duration of each waggle is closely correlated with the direction and distance from the flower patch being described. Mating behavior Within the honey bee colony, a queen bee typically mates with 10 or more males. 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. Apart from the queen bee, the only other sexual members of the society are the male drones, whose only function is to mate with the queen, after which they will die. 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 non-species drones present increases. 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. Scout bees search for suitable cavities in which to construct the swarm's home. 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. Absconding behavior 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==
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. 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. 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. ==Kin selection==
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 0.75 and those of different fathers will have a genetic relatedness of only 0.25. The females workers in the colony are related to the queen's sons by a genetic relatedness of 0.25. Such biasing results in the genes of some female worker bees being represented disproportionately in the virgin queens. 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. In other words, their fertility is controlled by queen signals. The queen honey bee informs workers of her presence by pheromones that she secretes from her mandibular glands. These signals are acquired by workers in close proximity to the queen and then spread to other workers in the colony, mainly by physical contact. In the presence of queen pheromone signals, the vast majority of workers refrain from activating their ovaries. Due to factors of genetic relatedness, an Apis cerana worker will 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. ==Interaction with other species==
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. Vespine wasps attempt to attack the honey bee quarry in an effort to gain provisions to aid in the development of their own offspring. though collection and use of feces in nest construction is well known in stingless bees. Pathogens and parasites affecting Apis cerana Microsporidia Apis cerana is the natural host to the microsporidian parasite Nosema ceranae, a serious pest of the western honey bee. When first discovered near Beijing, China, in 1994, it was originally thought that Nosema ceranae was restricted to Apis cerana in the East Asian region. 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. Sacbrood viruses (SBV) primarily affect the brood of the honey bee and causes larval death. Infected larvae fail to pupate, and ecdysial fluid aggregates around the integument, forming the "sac" for which the disease is named. Infected larvae change in color from pearly white to pale yellow, and shortly after death they dry out, forming a dark brown gondola-shaped scale. SBV may also affect the adult bee, but in this case obvious signs of disease are lacking. ==An invasive species in Australia==
An invasive species in Australia
Apis cerana was first detected in Australia in 2007. By 2012, it had spread across 500,000 hectares. The impacts of Apis cerana on the Australian environment are not well known due to limited research. However, according to Biosecurity Queensland (2103), the Apis cerana "is likely to compete for pollen and nectar with native birds, mammals and insects, and for nesting sites in tree crevices". There is a strong possibility that Apis cerana will also compete for resources with commercial honey bees and affect primary producers who rely on their pollination services. Control costs are also significant and amounted to at least up to 2011. Efforts to eradicate Apis cerana in Australia have failed. Although an eradication program commenced in 2007, a decision that it was not possible to eradicate Apis cerana was made in 2011. The bee is known as the Asian honey bee in Australia ==Genetic database==
Genetic database
As of 2015 the Biomodeling Laboratory at Seoul National University had constructed an Asian honey bee transcriptome database using an advanced sequencing technique. ==References==
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