Apis andreniformis | |
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Apis andreniformis, British Natural History Museum specimen | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Apidae |
Genus: | Apis |
Subgenus: | Micrapis |
Species: | A. andreniformis |
Binomial name | |
Apis andreniformis F. Smith, 1858 | |
Range of Apis andreniformis |
Apis andreniformis, or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia. [1]
A. andreniformis was the fifth honey bee species to be described of the seven known species of Apis . [1] Until recently, however, the actual identity of the species was poorly understood. It was not recognized as its own species, but was instead considered to be a part of the species Apis florea. [1] Recent studies have highlighted notable differences between the bees and have thus separated them into distinct species. [1]
Apis andreniformis is a part of the family Apidae, which includes honey, cuckoo, carpenter, digger, bumble, and stingless bees. [2] The genus Apis includes honey bees, the most common being Apis mellifera , otherwise known as the Western honey bee. A. andreniformis is most closely related to Apis florea, its sister species with which it is commonly seen in sympatric distribution throughout southeast Asia. [1]
A. andreniformis can be distinguished from other Apis species by noting their dark black coloration, making them the darkest of their genus. [3] Originally, it was thought that A. andreniformis was a part of the species A. florea, but recent studies have noted morphological differences that have separated the two. Some distinctions include: structural differences in the endophalli, a larger wing venation in A. andreniformis, and a longer basitarsal extension in A. florea. [4] Additionally, there are slight color variations between the two species. In typical A. andreniformis, its first two abdominal segments are black and its scutellum is reddish brown, while in A. florea, the first two abdominal segments are reddish brown and their scutellum is black, though there is some variation that makes color unreliable. [5] Another distinguishing factor is the presence of black hairs on the tibia of A. andreniformis, which are white in A. florea. [3]
Other differentiating characteristics include cubital indexes and proboscis length. A. andreniformis has an index of 6.37, while A. florea has one of 2.86. The proboscis of A. andreniformis has a length of 2.80 mm, while that of A. florea is 3.27 mm. [6]
Within the species, queens can typically be distinguished from workers and drones by their near entire black coloration. [3]
A. andreniformis nests are made of a single comb found hanging from small twigs [7] in quiet forests, generally in darker areas where there is 25 to 30% of normal sunlight. This type of nest is called an open-air nest. [8] They are commonly found hanging in small trees, shrubs, or bushes are usually hidden behind leaves or branches to avoid detection. [7] They are usually built between 1 and 15 meters from the ground, though the average altitude is 2.5 m. The honeycomb typically ranges from 70 to 90 mm in size. This nest is distinct from other Apis species, like A. mellifera , who build their nests inside of cavities. [8] This open-air structure—found also in A. florea, A. dorsata , and A. laboriosa —along with a relatively flat line of nectar cells along the top, creates a plateau above the nest that can be used as a stage for their communication method known as the waggle dance. [8]
While creating the nest, plant resin is placed along the supporting branch and around the edges of the nest. This acts as a barrier against small insects, like ants, that may try to enter the nest. [9] The major location of honey storage can be found in the area above and surrounding the branch. [7] The entire area below the honey storage and branch is the brood area, where larva development occurs. [7] Along the top of the brood area is the location of pollen storage. [7] Drone development occurs in the cells toward the bottom of the nest, while queen cells can be found protruding vertically. [7]
A. andreniformis is found in southeast Asia, specifically southern China, India, Burma, Laos, Vietnam, Thailand, Malaysia, Indonesia, and the Philippines. [1] [3] They are commonly found in sympatric distribution with A. florea. [5] Although they are sympatrically distributed, it is uncommon to have nests of different species in the same tree or bush. [9] Each species tends to be found closer to nests of its own species rather than its sister species. [9] A. andreniformis is considered a lowland species because they are most commonly found in elevations below 1,000 m, [9] although they may migrate to higher elevations during rainy seasons. [10] Similarly, they are found in tropical and subtropical regions, while cavity-dwelling honey bees can be found in colder climates. [1]
Queens are not genetically determined so any young female larva can become either a worker or a queen. [9] In order to create a queen cell, royal jelly must be fed to the female larva. [9] In cases of an unexpected loss of a queen, royal jelly is fed to a female worker larva to create a new emergency queen. [9] Although emergency queen rearing is possible, most often, the loss of a queen results in the dispersal of a colony. [9] If a queen becomes separated from her hive, the workers will leave the nest to search for her. Finding and joining a queen is an innate response for workers. The workers' strong attraction to queens is seen when a queen is lost, after a hive disruption by a predator, and during colony migration. [11]
In an Apis andrenformis queenright colony, worker policing occurs, which prevents workers from laying their own eggs and helps maintain the dominance hierarchy. [12] After the loss of a queen, workers will first try to rear a new queen using royal jelly. [9] If this fails, worker policing will decrease and workers will activate their ovaries in order to rear more drones before the colony dies. [12] It is also possible for queenless colonies to adopt a queen from a related species, specifically Apis florea. [11] This may suggest that worker bees of A. andreniformis are attracted to queen bee pheromones, even those belonging to other species in the same genera. [11] Once inside the colony of A. florea, the non-natal workers of A. andreniformis may attempt to lay their own eggs in order to continue propagating their species, an action known as reproductive parasitism. [12] In a study conducted by Sitthipong Wongvilas et al., they found that most eggs laid by the non-natal A. andreniformis were policed by the queenright workers of the A. florea colony. [12] Although they adopted A. andreniformis workers in the hive, A. florea workers policed non-natal larvae, thereby preventing interspecies parasitism. [12] Queenright colonies of A. andreniformis have also been observed to adopt queenless workers of A. florea as well, but their policing on non-natal larvae has not yet been studied. [12]
Unlike cavity-dwelling honey bees whose queen has a distinct chemical signal from that of the worker bees, A. andreniformis queens have similar chemical signals as their workers. [13] Chemical signals secreted from the mandibular gland in A. andreniformis are not caste-determining like it is in other honey bees. [13] As stated previously, the presence of royal jelly on young female larva produced the queen bee. [9] Drones, or male bees, are not used for pollination or honey production, but are instead used only to mate with the queen.
Different types of honey bees may use different types of dances to communicate with their hive. Most cavity-dwelling species use vertical waggle dances, while open-air nesters do not perform a gravity oriented waggle dance and instead perform a horizontal dance. [8] The shape of the nest creates a platform above the nest that can be used as a stage for communication. [8] The dance is a straight run pointing directly to the source of pollen or nectar that the forager has visited. Since the dance of other Apis species is vertical, it is not actually directed towards the food source, as it is in A. andreniformis.
Queens of A. andreniformis commonly engage in polyandry, where the queen will mate with multiple drones, usually about 10-20 times in total. [14] Due to the large amount of mating, queens must expel any excess semen. [14] Excessive mating puts females at increased risk to predation since it must occur during flight and outside the nest in the open air. She is also at risk for sexually transmitted diseases and injury from unexpected inclement weather. [14] Some Apis males put a "mating sign" in the sting chamber of the queen that she is unable to remove. This prevents her from avoiding unwanted copulation with other drones. In contrast, this sign is not found in A. andreniformis, suggesting that queens have control over the number of mates they copulate with. [14] Although there is a lot of risk to the queen, benefits may arise from the increased genetic diversity within the colony. Genetic diversity can lead to increased resistance to disease and illnesses. [14]
In studies, A. andreniformis has shown a lack of recognition for its own species and nestmates. [10] This has been shown in studies where queenless colonies of A. florea have joined the colony of A. andreniformis without facing aggression upon their initial arrival. [10] [11] Similarly, queenless colonies of A. andreniformis have been seen to join A. florea colonies, but in these cases, any A. andreniformis larvae are usually destroyed by the host colony, preventing parasitism from the foreign species. [11] Worker bees of A. florea have complete reproductive dominance over A. Andreniformis in a queenless nest because they have recognition and kin selection, while A. Andreniformis does not. However, when a queen is present, worker bees do not reproduce and parasitism is turned off. [12]
The main parasites of both A. andreniformis and A. florea belong to genus Euvarroa . However, A. andreniformis is attacked by the species Euvarroa wongsirii , while Euvarroa sinhai preys on A. florea, although Euvarroa sinhai have been found in hive debri of A. mellifera colonies it has not yet been confirmed to parasitize on them. The two species of Euvarroa have morphological and biological differences: while E. wongsirii has a triangular body shape and a length of 47 to 54 micrometres, E. sinhai has a more circular shape and a length of 39 to 40 micrometres. [15]
Honey bees, as a whole, tend to provide many useful products for human consumption. For A. andreniformis specifically, some commercial products include royal jelly, wax, honey, and bee venoms. [3] Additionally, they are important for the pollination of flowers and plants. [3]
A honey bee is a eusocial flying insect within the genus Apis of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmopolitan distribution of honey bees, introducing multiple subspecies into South America, North America, and Australia.
The honey bee life cycle, here referring exclusively to the domesticated Western honey bee, depends greatly on their social structure.
A queen bee is typically an adult, mated female (gyne) that lives in a colony or hive of honey bees. With fully developed reproductive organs, the queen is usually the mother of most, if not all, of the bees in the beehive. Queens are developed from larvae selected by worker bees and specially fed in order to become sexually mature. There is normally only one adult, mated queen in a hive, in which case the bees will usually follow and fiercely protect her.
A worker bee is any female bee that lacks the reproductive capacity of the colony's queen bee and carries out the majority of tasks needed for the functioning of the hive. While worker bees are present in all eusocial bee species, the term is rarely used for bees other than honey bees, particularly the European honey bee. Worker bees of this variety are responsible for approximately 80% of the world's crop pollination services.
Varroa destructor, the Varroa mite, is an external parasitic mite that attacks and feeds on honey bees and is one of the most damaging honey bee pests in the world. A significant mite infestation leads to the death of a honey bee colony, usually in the late autumn through early spring. Without management for Varroa mite, honey bee colonies typically collapse within 2 to 3 years in temperate climates. These mites can infest Apis mellifera, the western honey bee, and Apis cerana, the Asian honey bee. Due to very similar physical characteristics, this species was thought to be the closely related Varroa jacobsoni prior to 2000, but they were found to be two separate species after DNA analysis.
The dwarf honey bee, Apis florea, is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, Apis andreniformis. First identified in the late 18th century, Apis florea is unique for its morphology, foraging behavior and defensive mechanisms like making a piping noise. Apis florea have open nests and small colonies, which makes them more susceptible to predation than cavity nesters with large numbers of defensive workers. These honey bees are important pollinators and therefore commodified in countries like Cambodia.
Apis koschevnikovi, Koschevnikov's honey bee, is a species of honey bee which inhabits Malaysian and Indonesian Borneo, where it lives sympatrically with other honey bee species such as Apis cerana.
Apis nigrocincta is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of the tribe Apini.
Apis dorsata, the rock bee or giant honey bee, is a honey bee of South and Southeast Asia. They are typically around 17–20 mm (0.7–0.8 in) long and nests are mainly built in exposed places far off the ground, like on tree limbs, under cliff overhangs, and under buildings. These social bees are known for their aggressive defense strategies and vicious behavior when disturbed. Though not domesticated, indigenous peoples have traditionally used this species as a source of honey and beeswax, a practice known as honey hunting.
The East African lowland honey bee is a subspecies of the western honey bee. It is native to central, southern and eastern Africa, though at the southern extreme it is replaced by the Cape honey bee. This subspecies has been determined to constitute one part of the ancestry of the Africanized bees spreading through North and South America.
The western honey bee or European honey bee is the most common of the 7–12 species of honey bees worldwide. The genus name Apis is Latin for "bee", and mellifera is the Latin for "honey-bearing" or "honey carrying", referring to the species' production of honey.
Eusociality, the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform at least one behavior characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.
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.
Bumblebees, like the honeybee collect nectar and pollen from flowers and store them for food. Many individuals must be recruited to forage for food to provide for the hive. Some bee species have highly developed ways of communicating with each other about the location and quality of food resources ranging from physical to chemical displays.
Worker policing is a behavior seen in colonies of social hymenopterans whereby worker females eat or remove eggs that have been laid by other workers rather than those laid by a queen. Worker policing ensures that the offspring of the queen will predominate in the group. In certain species of bees, ants and wasps, workers or the queen may also act aggressively towards fertile workers. Worker policing has been suggested as a form of coercion to promote the evolution of altruistic behavior in eusocial insect societies.
Queen mandibular pheromone, or QMP, is a honey bee pheromone produced by the queen and fed to her attendants who share it with the rest of the colony to give the colony the sense of belonging to the queen. Newly emerged queens produce very little QMP. By the sixth day they are producing enough to attract drones for mating. A laying queen makes twice that amount. Lack of QMP seems to attract robber bees. A study of foraging worker bees has suggested that foraging bees are not attracted to QMP.
Nannotrigona testaceicornis is a eusocial stingless bee species of the order Hymenoptera and the genus Nannotrigona. Its local common name is abelhas iraí. This species has a large geographic distribution and occupies different biomes, including urban areas, around Neotropical America. The bees of this species nest in trees or artificial cavities because of this broad distribution. N. testaceicornis is important for agriculture because it will pollinate a vast number of plant species year round.
Scaptotrigona mexicana is a species of stingless bee that lives throughout Mexico and is part of the Meliponini tribe. This species is sometimes termed "Pisil Nekmej" and is extensively studied for its medicinal purposes. This species is considered common and abundant throughout Mexico and it has been noted to thrive in tropical environments.
Melipona scutellaris is a eusocial stingless bee species of the order Hymenoptera and the genus Melipona. It is considered to be the reared Melipona species with the largest distribution in the North and Northeast regions of Brazil, with records from Rio Grande do Norte down to Bahia. Its common name, Uruçu, comes from the Tupi "eiru su", which in this indigenous language means "big bee". Their honey is highly desirable and the materials they create for nests have been proven to be a promising source of antibiofilm agents and to present selectivity against human cancer cell lines at low concentrations compared to normal cells.
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