Apis of the world

author(s) : Pauly A.
Geographic zone: World.
Citation: Pauly A. 2015. The species of the genus Apis Linnaeus.
http://www.atlashymenoptera.net/page.aspx?ID=238

This is an attempt to present the systematics and biogeography of the species and subspecies in the genus Apis, with a list of most recently published papers.

Apis Linné

Africa - Asia - European bees
Apis (Apis) spp.
Middle-sized species with multiple combs in cavities; dances on vertical surfaces of combs in the dark (Michener, 2000). Four species: Apis mellifera, A. cerana, A. koschevnikovi and A. nigrocincta. Apis nuluensis is recognized at species level by some authors.
Apis (Apis) cerana Fabricius 1793
The Asiatic honeybee.

In the wild, A. cerana prefer to nest in small spaces, such as hollowed out tree trunks. They are similar in size or somewhat smaller than Apis mellifera and have smaller comb cells as well. Their honey yield is smaller, because they form smaller colonies. Like the Western honey bee, they are sometimes domesticated and used in apiculture, mostly in wooden boxes with fixed frames.
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Apis (Apis) cerana cerana Fabricius 1793
Distribution: as far west as Afghanistan and south along the central deserts and mountain ranges, across most of central and southern China, along the eastern edge of Asia up to Korea and Ussuria, and south to Northern Vietnam (Engel, 1999).

Hind wing venation
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Apis (Apis) cerana heimifeng Engel 1999
Distribution: Central China, in relatively high elevation areas (northern Sichuan Province, southwestern Gansu Province, and estern Qinghai Province).
Diagnosis: Distinguishable from typical A. cerana cerana in China by the dark brown to black scutellum and T3-T4 (these are yellow in A. cerana cerana). In this respect it resembles the Tibetan Honey Bee, A. cerana skorikovi, from which it differs in larger body size (body lenght 12.5-13.7 mm; forewing lenght 8.8-9.3 mm).
Source: Engel, 1999.
Apis (Apis) cerana indica Fabricius 1798
Distribution: India.
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Apis (Apis) cerana japonica Radoszkowski 1887
Distribution: Japan
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Apis (Apis) cerana javana Enderlein 1906
Distribution: Java to Timor.
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Apis (Apis) cerana johni Skorikov 1929
Distribution: Sumatra
Apis (Apis) cerana nuluensis Tingek 1996
Distribution: mountains of Sabah
Otis 1996.
Apis (Apis) cerana philippina Skorikov 1929
Philippines
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Apis (Apis) cerana skorikovi Engel 1999
Apis cerana skorikovi Maa 1944, nomen nudum.
Apis cerana, Himalaya race, Ruttner 1988.
Apis cerana, Xizang race, Peng et al. 1989.
Apis cerana himalaya Smith 1991, nomen nudum.

Distribution: along the Himalayan uplift at elevations of 1900-4000 m.
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Apis (Apis) koschevnikovi Enderlein 1906
The Sundaland Honey Bee

Apis mellifica indica var koschevnikovi Buttel-reepen 1906. Unavailable because used infrasubspecifically (I.C.Z.N. 1985: Arts 10c, 23j, and 50c).
Apis indica variety koschevnikovi Enderlein 1906. First available usage.
Apis (Sigmatapis) vechti Maa 1953
Apis (Sigmatapis) vechti linda Maa 1953
Apis mellifera adansoni koschevnikowi Goetze 1964. Unjustified emendation.

Distribution: Malay Peninsula, Borneo, Brunei, Java, Sabah, Sarawak, and Sumatra (Engel, 1999).

Diagnose. This species is sympatric to A. cerana. It can be separated by it's black colour, and the different endophallus and hind legs of the drone (see Tingek et al. 1988).
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Apis (Apis) mellifera Linnaeus 1758
Honey bee
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Apis (Apis) mellifera adami Ruttner 1975
Distribution: island of Crete.

The illustrated specimen is from Vrakhas-H. Georgios, 250 m, , 29.IV.1931 (RBINS, Brussels).

Following Harizanis & Bouga (2003), it seems that there are today no pure populations of Apis m. adami in Crete.
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Apis (Apis) mellifera adansonii Latreille 1804
Distribution: Forests of Central and West Africa.
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Apis (Apis) mellifera anatoliaca Maa 1953
Distribution: Turkey
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Apis (Apis) mellifera artemisia Engel 1999
Apis mellifera mellifera natio tesquorum Skorikov 1929, unavailable (I.C.Z.N. 1985: Arts. 10c, 23j, 50c)
Apis mellifera acervorum Skorikov 1929. Preoccupied (nec Linnaeus 1758)

Distribution: central Russian Steppes.
Apis (Apis) mellifera capensis Eschscholtz 1822
Distribution: Cape region of South Africa.

Comment: This subspecies is biologically distinctive for its ability to undergo parthenogenetic reproduction when deprived of a queen (Jack 1916, Anderson 1963, Ruttner 1977).
Apis (Apis) mellifera carnica Pollmann 1879
Distribution: European race occuring south of the Alps, east into northern Italy, and west into Yugoslavia and Romania.
Apis (Apis) mellifera caucasia Pollmann 1889
Apis mellifida (sic!) variety caucasica Pollmann 1889.
Apis mellifera caucasica Gorbachev 1916. Unjustified emendation.

Distribution: Caucasus Mountains.
The "grey" Caucasian honey bee.
Apis (Apis) mellifera cecropia Kiesenwetter 1860
The Greek Honey Bee

Distribution: Occurs throughout most of Greece and surrounding Aegean islands.

Reference: Bouga & Hatjina (2005).
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Apis (Apis) mellifera cypria Pollmann 1879
The Cyprian Honey Bee

Distribution: island of Cyprus
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Apis (Apis) mellifera iberiensis Engel 1999
Apis mellifera intermissa iberica Goetze 1964. Infrasubspecific name proposed after 1960 and therefore unavailable (I.C.Z.N. 1985: Art. 1b); preoccupied (nec. Skorikov 1929).
Apis mellifera iberica Ruttner 1988: 236. Corrected authorship (first available usage); preoccupied (nec Skorikov 1929).

Distribution: natively occurs on the Iberian Peninsula in Europe.
Two ecotypes of A. mellifera iberiensis occur in Spain as shown by Santiago et al. (1986) as well as two mitochondrial types as demonstrated by Smith et al. (1991) and Smith and Glenn (1995). Neotype designated by Engel from Cordoba populations sampled by Smith and Glenn (1995: their colony 1-1).
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Apis (Apis) mellifera intermissa Maa 1953
The Tellian Honey Bee

The name intermissa as used by Buttel-reepen (1906) is unavailable. It was first made available by Maa (1953).

Distribution. This race has a tight distribution along the northern coast of Africa as far west as Morocco, into Tunisia in the east, but bordered by the Atlas range in the south.
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Apis (Apis) mellifera jemenitica Ruttner 1976
Reinstated name by Engel 1999, The Arabian or Nubian Honey Bee.

Apis mellifera nubica Ruttner 1976.
Apis mellifera yemenitica Ruttner 1988. Unnecessary emendation.
Apis mellifera sudanensis Ruttner 1988. Nomen nudum.

No type designed for this subspecies (Engel 1999). Ruttners original material in Oberursel.

Distribution. Sahelian countries, from Senegal to Arabia.

Geographical distribution: Al-Ghamdi et al. 2013.
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Apis (Apis) mellifera lamarckii Cockerell 1906
The Egyptian Honey Bee.

Distribution: This honey bee race occurs in a narrow range along the Egyptian Nile Valley.

Remark: replacement name for Apis fasciata Latreille 1804, preoccupied (nec Linnaeus 1767, nec Scopoli 1770).
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Apis (Apis) mellifera ligustica Spinola 1806
The Italian Honey Bee

Distribution: occurs along the Italian Peninsula. it has been commercially trasnported throughout the world.
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Apis (Apis) mellifera litorea Smith 1961
Distribution: along the eastern coast of tropical Africa occuring from Kenya (perhaps even the southern-most portions of Somalia) south to Mozambique.

No type appear to have been designated for this species (Engel 1999).
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Apis (Apis) mellifera macedonica Ruttner 1988
The Macedonian Honey Bee

Distribution: southern Romania, east to Yugoslavia, and south to northern Greece (Engel 1999).

No type has been designated fro this race (Engel 1999). Original material in Ruttner collection.
Apis (Apis) mellifera meda Skorikov 199
The Median Honey Bee

Distribution: Iran, Iracq, southeastern turkey and northern Syria.
Apis (Apis) mellifera mellifera Linnaeus
The Western or European Honey Bee

Distribution: originally ranged throughout central Europe north of the Alps, as far south as southern Sweden in the north, central russia in the east, and on the British Isles. In the Ukraine there is a transition over the steppe region to A. mellifera sossimai (Engel 1999).
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Apis (Apis) mellifera monticola Smith 1961
The East African Black Mountain Honey Bee

Distribution: Mountains from Kenya and Tanzania. Described from Kilimandjaro but no type appears to have been designated (Engel 1999).

Mountain honeybees of Africa: see Hepburn et al. (2000)
Voir Meixner et al. 1989.
Phenotypic plasticity: Gruber et al. 2013.
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Apis (Apis) mellifera remipes Gerstaecker 1862
The Yellow Armenian Honey Bee
= Apis mellifera armeniaca Skorikov, 1929

Name reinstated by Engel 1999.

Distribution: This race occurs in Armenia and may be the same as A. mellifera anatoliaca, in which case Gerstaecker's name has priority.
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Apis (Apis) mellifera ruttneri Sheppard, Arias, Grech & Meixner 1997
Distribution: island of Malta in the Mediterranean Sea.
Apis (Apis) mellifera sahariensis Baldensperger 1932
The Saharan Honey Bee

Distribution: southern side of the Atlas range in North Africa.

No type designated (Engel 1999).

Possibly a synonym of A. m. lamarckii from Egypt.
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Apis (Apis) mellifera scutellata Lepeletier de Saint Fargeau 1836
The African Honey Bee.

Range from South Africa and northward along eastern half of the continent.

This species was originally described from "Caffraria" (in South Africa) and distinguished from Apis caffra (Lepeletier de saint Fargeau 1836) (= A. m. capensis) by its yellow scutellum.

Remark: tis is the race introduced into Brazil in the 1950's, it is known as the "Africanized Honey Bee" or "Killer Bee".

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Apis (Apis) mellifera siciliana Grassi 1881
The Sicilian Honey Bee

Name reinstated by Engel 1999

Apis siciliana Grassi 1881
Apis mellifica variety siziliana Buttel-Reepen 1906. Unjustified emendation.
Apis sicula Montagano 1911.

Distribution: island of Sicily in the Mediterranean Sea.
Apis (Apis) mellifera simensis Meixner et al. 2011
Distribution: High plates of Ethiopia
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Apis (Apis) mellifera sossimai Engel 1999
The Ukarinian Honey Bee

Apis cerifera Gerstaecker 1862. Preoccupied (nec scopoli 1770).

Distribution: Ukraine, Crimea excepted. Northern regions of the Caucasus Mountains.
Apis (Apis) mellifera syriaca Skorikov 1929
The Syrian Honey Bee or the Palestine Honey bee.

Apis mellifera syriaca as used by Buttel-Reepen (1906) is unavailable. Corrected authorship by Skorikov 1929 (Engel 1999).

Distribution: eastern shores of the Mediterranean Sea; north from syria to the Negev Desert in the south.
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Apis (Apis) mellifera taurica Alpatov 1935
The Crimean Honey Bee

Distribution: along the north-central shores of the Black Sea; in the Crimea.
Apis (Apis) mellifera unicolor Latreille 1804
The Malagasy Honey Bee.

Distribution: Madagascar.

Brooks & Michener, 1988.
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Apis (Apis) nigrocincta Smith 1861
The Sulawesian Honey Bee

Apis nigrocincta Smith 1861
Apis mellifica indica var picea Buttel-Reepen 1906
Apis (Sigmatapis) nigrocincta marginella Maa 1953

Literature: Hadisoesilo et al. 1995, Hadisoesilo and Otis 1996, 1998, Hadisoesilo 1997.

Distribution: endemic to Sulawezi.

Diagnose. The drone cells of A. nigrocincta are soft and lack pores, unlike those of A. cerana which are always herdened and have pores. Drones of A. cerana always took their mating flights about 2 hr earlier than do drones of A. nigrocincta, with little overlap in the distributions of flight times. The distributions of A. cerana and A. nigrocincta are largely allopatric, with narrow zone of overlap. In Central Sulawezi, the abrupt transition corresponds to the restriction of A. nigrocincta to forests and of A. cerana to disturbed agricultural areas. The male genitalia of both species are similar. (source: Hadisoesilo 1997).
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Apis (Megapis) spp
Large species with single exposed combs; dances on vertical curtains of bees or on comb.
Apis (Megapis) dorsata binghami Cockerell 1906
Distribution: Sulawezi
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Apis (Megapis) dorsata breviligula (Maa 1953)
Distribution: Philippines.
The species status is oft argued based on nest site characteristics as reported by Morse and Laigo (1969) and Starr et al. (1987)
Molecular phylogeny: see Lo et al. (2010)
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Apis (Megapis) dorsata dorsata Fabricius 1793
Distribution: From India, east to the coast of Vietnam, and into the southeast Asian islands where it is sometimes replaced by other subspecies (Engel, 1999).

Nests are mainly built in exposed places far off the ground, on tree limbs and under cliff overhangs, and sometimes on buildings. Apis dorsata is a defensive bee and has never been domesticated (as it does not use enclosed cavities for nesting). Each colony consists of a single vertical comb (sometimes approaching a square metre) suspended from above, and the comb is typically covered by a dense mass of bees in several layers. When disturbed, the workers may exhibit a defensive behavior known as defense waving.
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Apis (Megapis) dorsata laboriosa Smith 1871
Distribution: At high altitudes (1200 to 4000 m) on the slopes of the mountains from northernmost India to the northernmost boundary of Laos (Engel, 1999).
Literature: Otis (1996), Lian-Fei Cao et al. (2012)
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Apis (Micrapis) spp.
Small species with single exposed combs; dances on expanded horizontal base of comb. Two species.
Apis (Micrapis) andreniformis Smith 1858
Distribution: north to China (Yunnan), south into Malaysia, eastward on the mainland to the coast of Vietnam, and westward as far as north-western India along the eastern border of Nepal (Engel, 1999).

Biogeography: Hepburn & Radloff 2011.

The most reliable character to rapidly distinguish A. andreniformis from A. florea is the denser punctuation of the abdominal tergite 2.

The mistaken notion that abdominal tergites 1 and 2 of A. florea are reddish and
other segments at least partially reddish, while those of A. andreniformis are
uniformly black, still permeates the literature (Radloff, Hepburn & Engel in Hepburn & Radloff, 2011, Honeybees of Asia).
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Apis (Micrapis) florea Fabricius 1787
Distribution. From the eastern region of the Arabian Peninsula, through southern Iran and Iraq to Pakistan and India as far north as the Himalayan uplift. The species extends eastward as far as Vietnam, north to Yunnan Province, China, and south into Indonesia (Engel, 1999).

Jordan (Haddad et al. 2009)
Invasive in Africa: first discovered in Sudan (Lord & Nagi, 1987), now also in northern Ethiopia (Pauly & Hora 2013).

Morphometry: Hepburn et al. 2005.
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References

Al-Ghamdi A., Alsharhi M., Alatal Y. & Nuru A. 2012. Morphometric diversity of indigenous Honeybees, Apis mellifera (Linnaeus, 1758), in Saudi Arabia. Zoology in the Middle East, 57: 97-103.
Al-Ghamdi A., Nuru A., Khanbash M.S. & Smith D.R. 2013. Geographical distribution and population variation of Apis mellifera jemenitica Ruttner. Journal of Apicultural Research, 52(3): 124-133.
Alvarez-Suarez J.M. Editor. 2017. Bee Products - Chemical and Biological Properties. Springer. 306 pp.
Amssalu B., Nuru A., Radloff S.E. & Hepburn H.R. 2004. Multivariate morphometric analysis of honeybees (Apis mellifera) in the Ethiopian region. Apidologie, 35: 71-81.
Anderson R.H. 1963. The laying worker in the Cape honeybee, Apis mellifera capensis. Journal of Apicultural research, 2: 85-92.
Bouga M. & Hadjina F. 2005. Genetic variability in Greek Honey Bee (A. mellifera L.) populations using geometric morphometrics analysis. Proceedings of the Balkan scientific conference of Biology in Plovdiv (Bulgaria) from 19th till 21st of may 2005 (Eds B. Gruev, M. Nikolova & A. Donev), 2005 (p. 598-602).
Chantawannakul P., Williams G. & Neumann P. Eds. 2018. Asian Beekeeping in the 21st Century. Springer, 325 pp.
see also:https://doi.org/10.1007/978-981-10-8222-1
Engel M. 1999. The taxonomy of recent and fossil honey bees. Journal of Hymenoptera Research, 8: 165-196.
Gruber K., Schöning C., Otte M., Kinuthia W. & Hasselmann M. 2013. Distinct subspecies or phenotypic plasticity? Genetic and morphological differentiation of mountain honey bees in East Africa. Ecology and Evolution, 1-15.
Haddad N., Fuchs S., Hepburn H.R. & Radloff S.E. 2009. Apis florea in Jordan: source of the founder population. Apidologie, 40: 508-512.
Hadisoesilo S. & Otis G.W. 1996. Drone flight times confirmthe species status of Apis nigrocincta Smith, 1861, to be a species distinct from Apis cerana F., 1793, in Sulwaezi, Indonesia. Apidologie, 27: 361-369.
Harizanis P. & Bouga M. 2003. Genetic structure of the Bee from Crete Island (Greece). XXVIIIth International Apicultural Congress, Final Programme and Book of Abstracts, Ljubljana, Slovenia, p. 696.
Hepburn H.R. & Radloff S.E. 1998. Honeybees of Africa. Springer. 370 pp.
Hepburn H.R. & Radloff S.E. 2011. Biogeography of the dwarf honeybees, Apis andreniformis and Apis florea. Apidologie, 42: 293-300.
Hepburn H.R. & Radloff S.E. 2011. Honeybees of Asia. Springer, 600pp.
Hepburn H.R., Radloff S.E. & Oghiakhe S. 2000. Mountain honeybees of Africa. Apidologie 31: 205-221.
Hepburn H.R., Radloff S.E., Otis G.W., Fuchs S., Verma L.R., Ken T., Chaiyawong T., Tahmasebi G, Ebadi R. & Wongsiri S. 2005. Apis florea: morphometrics, classification and biogeography. Apidologie, 36: 359–376.
Jack R.W. 1916. Parthenogenesis amongst the workers of the Cape honey-bee: Mr.G.W. Onion's experiments. Transactions of the Entomological Society of London, 1916: 396-403.
Lian-Fei Cao, Huo-Qing Zheng, Xuan Chen, De-Fang Niu, Fu-Liang Hui & H R. Hepburn. 2012. Multivariate morphometric analyses of the giant honey bees, Apis dorsata F. and Apis laboriosa F. in China. Journal of Apicultural Research 51(3): 245-251.
Lo N., Gloag R.S., Anderson D.L. & Oldroyd B.P. 2010. A molecular phylogeny of the genus Apis suggests that the Giant Honey Bee of the Philippines, A. breviligula Maa, and the Plains Honey Bee of southern India, A. indica Fabricius, are valid species. Systematic Entomology (2010), 35, 226–233.
Lord W.G. & Nagi S.K. 1987. Apis florea discovered in Africa. Bee World, 68 (1): 39-40.
Maa T.C. 1953. An inquiry into the systematics of the tribus Apidini or honeybees (Hym.). Treubia, 21: 525-640.
Meixner M., Ruttner F., Koeniger N., Koeniger G. 1989. The mountain bees of the Kilimanjaro region and the relation to neighbouring bee populations. Apidologie, 20: 165-174.
Meixner M.D., Leta M.A., Koeniger N. & Fuchs S. 2011. The honey bees of Ethiopia represent a new subspecies of Apis mellifera – Apis mellifera simensis n. ssp. Apidologie, 42: 425-437.
Ming-Xian Yang. 2009. Studies on Mixed–Species Colonies of Honeybees, Apis cerana and Apis mellifera. Thesis submitted in fulfilment of the requirements For the Degree of Doctor of Philosophy At Rhodes University. 132pp.
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Oldroyd B.P., Chapman R.N.C., Thompson G.J. & Beekman M. 2006. Evidence for reproductive isolation between two colour morphs of cavity nesting honey bees (Apis) in south India. Insectes Sociaux, 53: 428-434.
Otis G.W. 1996. Distribution of recently recognized species of honey bees (Hymenoptera: Apidae; Apis). Journal of the Kansas Entomological Society, supplement 69, 311-333.
Pauly A. & Hora Z.A. 2013. Apini and Meliponini from Ethiopia (Hymenoptera: Apoidea: Apidae: Apinae) . Belgian Journal of Entomology, 16: 1-36.
Pinto M.A., Munoz I., Chavez-Galararza J., De la Rua P. 2012. The Atlantic side of the Iberian Peninsula: a hot-spot of novel African honey bee maternal diversity. Apidologie, 43: 663-673.
Rinderer T.E., Koeniger N., Tingek S., Mardan M. & Koeniger G. 1989. A morphological comparison of the cavity dwelling honeybees of Borneo Apis koschevnikovi (Buttel-Reepen, 1906)* and Apis cerana (Fabricius, 1793). Apidologie, 20: 405-411.
Ruttner F. 1977. The problem of the Cape bee (Apis mellifera capensis Escholtz): parthenogenesis, size of populations, evolution. Apidologie, 8: 281-294.
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Starr C.K., Schmidt P.J. & Schmidt J.O. 1987. Nest-site preferences of the giant honey bee, Apis dorsata (Hymenoptera: Apidae), in Borneo. Pan Pacific Entomologist, 63: 37-42.
Suwannapong G., Benbow M.E. & Nieh J.C. 2011. Biology of Thai Honeybees: natural history and threats . In: Bees: Biology, Threats and Colonies, Editor: Richard M. Florio, pp. 1-98.
Tingek S., Mardan M., Rinderer T.E., Koeniger N. & Koeniger G. 1988. Rediscovery of Apis vechti (Maa, 1953): The Saban Honey Bee. Apidologie, 19 (1): 97-102.