''Apis koschevnikovi'', Koschevnikov's honey bee, is a species of

honey bee A honey bee (also spelled honeybee) 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 ...

which inhabits

Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...

n and

Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...

Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...

, where it lives sympatrically with other honey bee species such as '' Apis cerana'' (specifically '' A. c. nuluensis''). The species was first described by Hugo Berthold von Buttel-Reepen, who dedicated it to Grigory Aleksandrovich Kozhevnikov (1866–1933), a pioneer of honey bee

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...

. This was an invalid designation, however, and the name was first formally made available by

Günther Enderlein Günther Enderlein (7 July 1872 – 11 August 1968) was a German zoologist, entomologist, microbiologist, researcher, physician for 60 years, and later a manufacturer of pharmaceutical products. Enderlein received international renown for his ...

that same year. Therefore, Buttel-Reepen is not the author of the name (following the

ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...

). The species was described again by Maa in 1953, this time with the name ''Apis vechti''. It was finally rediscovered by Tingek and his colleagues in 1988.

Taxonomy and phylogeny

''Apis koschevnikovi'' is of the family Apidae and genus ''Apis''. ''A. koschevnikovi'' is known as one of the "Red Bees" of Borneo, described in 1988. ''A. koschevnikovi'' appears together with '' A. cerana'' and '' A. mellifera'', two other cavity-nesting species, in three separate phylogenic clusters without overlapping. The

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

cluster analysis of ''A. koschevnikovi'' is found directly in contact with a cluster of ''A. cerana'' and distant from ''A. mellifera''.

Description and identification

''A. koschevnikovi'' is often referred to in the literature as the "red bee of Sabah"; however ''A. koschevnikovi'' is pale reddish in Sabah State,

, but a dark, coppery color in the

Malay Peninsula The Malay Peninsula is located in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area contains Peninsular Malaysia, Southern Tha ...

and

Sumatera Sumatra () is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 482,286.55 km2 (182,812 mi.2), including adjacent i ...

Physical appearance

The workers, the

queen Queen most commonly refers to: * Queen regnant, a female monarch of a kingdom * Queen consort, the wife of a reigning king * Queen (band), a British rock band Queen or QUEEN may also refer to: Monarchy * Queen dowager, the widow of a king * Q ...

, and the drones of ''A. koschevnikovi'' are all dark brown banded. The workers, however, have light orange abdominal bands while the queen and the drones have light brown abdominal bands. '' Ko'' is the gene responsible for the expression of body color type in ''A. koschevnikovi''. This gene is shown to be sex-linked, since experiments have shown that a cross between a brown, dark banded queen and a brown, dark banded drone results in orange, dark banded workers. ''A. koschevnikovi'' is very long-tongued (5.870 mm) and slender, with narrow tomenta. ''Apis koschevnikov''i is larger than its

sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...

'' A. cerana'', consistently being 10 to 15% larger linearly. ''Apis koschevnikovi'' also has two distinct characteristics which are species specific – its drones have a secondary sex characteristic of a hairy fringe on the margin of the tibia of the hind leg, and worker bee forewing venation shows a cubital index which is large and varied.

Distribution and habitat

The habitat of ''A. koschevnikovi'' is limited to the tropical evergreen forests of the

, and

Sumatra Sumatra () is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...

. They do not live in tropical evergreen rain forests which extend into

Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...

Myanmar Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and has ...

Cambodia Cambodia, officially the Kingdom of Cambodia, is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. It is bordered by Thailand to the northwest, Laos to the north, and Vietnam to the east, and has a coastline ...

, and

Vietnam Vietnam, officially the Socialist Republic of Vietnam (SRV), is a country at the eastern edge of mainland Southeast Asia, with an area of about and a population of over 100 million, making it the world's List of countries and depende ...

. This area is associated with a change from wet seasonal evergreen rain forests to mixed moist deciduous forests. Its altitudinal distributions extend from sea level to about 1600 meters. The range for ''A. koschevnikovi'' is diminishing because it is now either poorly represented or absent in the several areas where it had been previously located. This has been attributed to habitat changes resulting from

deforestation Deforestation or forest clearance is the removal and destruction of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. Ab ...

and the establishment of tea, oil palm, rubber, and coconut plantations. Not much is known about the nest structure of ''Apis koschevnikovi''. They live in small colonies making a few combs within small tree cavities in the

rain forest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...

Behavior

Mating behavior

The drones for ''Apis koschevnikovi'' have a specific mating time that is species-specific. Regardless of whether or not they were in a

conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...

or other species' colony, the drones will fly at their species specific mating time. The queens also follow their own internal clock. This allows for a reproductive isolation mechanism that is based on individual behavior of the sexual castes.

Costs and benefits of sociality

''Apis koschevnikovi'' have a very small colony size of a thousand bees or so. This small colony size allows them to survive in a rain forest habitat. Despite their small colony size, they are still able to harvest resources very quickly and reproduce at a fast pace during the general flowering period.

Queen acceptance of different species

There is a strong tendency for preference of

larvae in ''Apis koschevnikovi''. ''A. koschevnikovi'' is even more selective with regard to the acceptance of a young alien

. When too young of a queen of a different species is introduced into the colony of an ''Apis koschevnikovi'', the alien queen uses a buzzing sound in response to the aggression from the worker bees. This is successful at first until the aggressive behavior towards the queen increases with the age of the queen. After the third or fourth day of emergence, the worker queens mutilate and expel the alien queen from the colony. The current hypothesis for the observed worker bee aggression is thought to be from the wrong blend of queen

pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...

s released from the alien queen.

Interaction with other species

Diet

''Apis koschevnikovi'' is a

nectar-feeder. ''Apis koschevnikovi'' is a steady resident component of the

and visits flowers during the entire year. In Bornean primary forests, '' Apis dorsata'' and ''A. koschevnikovi'' are the only honeybees that appear frequently at flowering canopy trees or at baits. At the Lambir Hills National Park in

, these two honeybees visited 29 flower species, making them visitors of 10% of the plant species and 22% of the plant families of the park.

Defense of resources

Even though ''Apis koschevnikovi'' and '' Apis dorsata'' share mostly the same rain forest habitat and overlap spatially in using trees at all heights in the canopy, they are still able to coexist. They differ in size and tongue length, which helps separate resource use. Only half of the resources that ''Apis koschevnikovi'' uses are shared with ''Apis dorsata''. However, in contrast to normal feeding behavior, ''Apis koschevnikovi'' and '' Apis dorsata'' fight at artificial feeding stations. They displayed a displaced nest-defense behavior with grappling and attempted stinging. Honey bees do not interact well with competing foragers when near a rich resource. ''A. koschevnikovi'' has even shown aggression against

foragers when competing for a resource. Honey bees tend to dominate floral patches as individual colonies because of their competitive nature.

Parasites

''A. koschevnikovi'' hosts a unique species of the honey bee parasitic mite genus '' Varroa'', named '' Varroa rindereri''. Although this

parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...

species is quite similar to '' Varroa jacobsoni'' it is perfectly differentiable. ''V. rindereri'' is larger (1 180 x 1 698 micrometers). ''V.rindereri'' also has a fewer number of

seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...

e and pores on the sternal shield. It has a long and wide-looped peritreme, and the trochanter of the palpus lacks a seta. When pupae were removed from their cells, ''V. rindereri'' remained inside the cells. It has only been reported in colonies of ''A. koschevnikovi'' in Borneo and seems to be specific to that species, as it has yet to be observed crossing over to colonies of '' A. cerana'', even when they live in the same

apiary An apiary (also known as a bee yard) is a location where Beehive (beekeeping), beehives of honey bees are kept. Apiaries come in many sizes and can be rural or urban depending on the honey production operation. Furthermore, an apiary may refer to ...

Interspecies broods

When a brood of ''A. cerana'' drones are hatched in a colony of ''A. koschevnikovi,'' they are fully taken care of and raised to maturity. Likewise when ''A. koschevnikovi'' drones are hatched in a colony of ''A. cerana'' they are completely integrated into the society of their host. However, it has been shown that despite the crossfostering of drones, each species will not change its mating time and flight habits, even if it is raised as an alien drone.

References

External links

New haplotypes for the non-coding region of mitochondrial DNA in cavity-nesting honey bees ''Apis koschevnikovi'' and ''Apis nuluensis''

Database of the mitochondrial DNA of ''Apis koschevnikovi''

Crossfostered drones of ''Apis cerana'' (Fabricius, 1793) and ''Apis koschevnikovi'' (Buttel-Reepen, 1906) fly at their species specific mating times
{{Taxonbar, from=Q728308 koschevnikovi Hymenoptera of Asia Insects of Borneo Insects described in 1906