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TREUBIA
A JOURNAL ON ZOOLOGY
OF THE INDO-AUSTRALIAN ARCHIPELAGO

Vol. 39, pp. 1-85 December 2012

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ISSN : 0082 - 6340
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TREUBIA
A JOURNAL ON ZOOLOGY OF THE INDO-AUSTRALIAN ARCHIPELAGO
Vol. 39, pp. 1-85, December 2012

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 Treubia 2012, 39: 41-49

THE HONEY BEES OF INDONESIA

(HYMENOPTERA: APIDAE)
Michael S. Engel

Division of Entomology, Natural History Museum, and

Department of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140,
University of Kansas, Lawrence, Kansas 66045, USA
email: [email protected]

ABSTRACT
A brief account is provided for the native and introduced species of honey
bees (Apini: Apis L.) occurring across Indonesia. Keys to the tribes of
corbiculate bees and the species of the genus Apis are provided to aid
melittologists and apiculturists working in the country.
Key words: Apoidea, Anthophila, Apinae, honey bees, Apis, identification keys

INTRODUCTION
Among the diversity of around 20,000 species of bees worldwide,
the honey bees (genus Apis Linnaeus) are the most heavily researched and
managed, focusing the efforts of thousands of researchers and tens of
thousands of apiculturists. However, in terms of bee diversity, Apis
accounts for only seven living species (Engel 1999) or 0.035% of that
remarkable diversification. Honey bees are one of four living tribes of
corbiculate apine bees. The lineage of corbiculate bees comprises the
orchid bees (Euglossini Latreille), bumble bees (Bombini Latreille),
stingless bees (Meliponini Lepeletier de Saint Fargeau), and the honey
bees (Apini Latreille) in the modern fauna (Engel 2001a, 2005, Michener
2007), as well as the extinct tribes Electrobombini Engel, Electrapini
Engel, and Melikertini Engel (Engel 2001a, 2005), the latter representing
the presumed sister group of the meliponines (Engel 2001a, 2001b). All
of these lineages are highly eusocial with the exceptions of the primitively
eusocial Bombini and Electrobombini, and the communal or solitary
Euglossini.
While today the honey bees are restricted to the Old World, and
with only a single species occurring in Europe (Apis mellifera Linnaeus)
and considerably reduced diversity in Africa (A. mellifera and A. florea
Fabricius), during the relatively recent geological past the genus was
much more widespread and included Western North America (Engel et al.
2009), and even regions such as Europe had multiple species and
exhibited dramatically greater morphological disparity (e.g., Kotthoff et

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 Michael S. Engel: The honey bees of Indonesia (Hymenoptera: Apidae)

al. 2011). Also, lineages which today are more restricted had greater
geographic distributions (e.g., Megapis species in Iki Island of the Korea
Strait: Engel 2006). Indeed, during the Oligocene and Miocene, the area
with greatest Apis diversity was Europe (Engel 1998, Kotthoff et al. 2011).
It is within Southeast Asia that honey bees today exhibit their greatest
diversity both at and below the species level. Indeed, at times this diversity
has seemed overwhelming and Maa (1953) recognized as many as 24
species within the genus, although more modern treatments have paired
this plethora considerably (e.g., Engel 1999, Radloff et al. 2011, Ruttner
1988). Surprisingly, despite the remarkably intense scrutiny that has been
applied to Apis diversity in Asia, few truly systematic works exist and
world or regional keys have been lacking or difficult to access by most
(e.g., Engel 2001c, 2002, Malaipan 1972). Maa’s (1953) keys remain the
only dichotomous means of recognizing the world’s taxonomic units for
honey bees, although revised treatments are underway (Engel in prep.).
Herein I provide an identification key to those species of honey bees
known to occur, both natively as well as from introduced species, across
Indonesia. Indonesia harbors natively five of the seven known species and
a remarkable breadth of variation within those species. Accordingly, I
have tailored the colors used in the key to reflect the ranges observed
within Indonesia and provided brief notes on the occurrence of those
species within the country, all as an aid for apiculturists working in the
country. The introduced Western Honey Bee (A. mellifera) is included so
as to aids its differentiation from the native species. Introduction of A.
mellifera has been attempted several times over the last 150 years but only
intensified during the last 20–25 years mostly as the result of commercial
efforts. Fortunately, many of these have failed but Indonesian apiculturists
should be aware of A. mellifera and encouraged to continue to cultivate,
even on larger industrial scales, their native species over this European
form. In addition, I include a key to the three living tribes of corbiculate
bees similarly known to occur in the region and in the hope of stimulating
study of these groups by local researchers. Such materials are being
published in advance and coordination of ongoing work related to the
world diversity of honey bees (Engel in prep.). Morphological terminology
used throughout is based on the more standard systems as applied to all
bees (and, in most cases, to all Hymenoptera) (Engel 2001a, Michener
2007), rather than the more regional or specialised terms sometimes
employed by apiculturists but which in some cases are erroneous or
misleading. The classification of species adopted is that of Engel (1999),
although some researchers prefer to elevate regionally distinctive forms to
specific rank despite its concomitant implications for the circumscription
of the related forms (e.g., refer to Radloff et al. 2011). Recent and relevant

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 Treubia 2012, 39: 41-49

biological accounts of the various honey bees may be found in Oldroyd &
Wongsiri (2006), Koeniger et al. (2010), and Hepburn & Radloff (2011),
as well as references cited in these works.
Key to the Tribes of Corbiculate Bees in Indonesia
Within Indonesia, three of the four living tribes of corbiculate bees are
known to occur. Honey bees (Apini) and stingless bees (Meliponini)
occur throughout the country, while only five species of bumble bees
(Bombini) are known from the western parts of Indonesia (Sumatran
region).
1. Jugal lobe of hind wing present; metatibial spurs absent; arolia
present; outer grooves of mandible absent ......................................... 2
—. Jugal lobe of hind wing absent; metatibial spurs present; arolia absent
or considerably reduced and vestigial; outer grooves of mandible
present (bumble bees) ............................................................ Bombini
2. Forewing with reduced distal venation, marginal cell frequently open
at apex; pretarsal claws simple; penicillum present in worker; auricle
absent; sting reduced (stingless bees) ..................................Meliponini
—. Forewing with complete distal venation, marginal cell long and
completely bordered by strong tubular veins; pretarsal claws cleft;
penicillum absent in worker; auricle present; sting well developed
and barbed (honey bees) ............................................................. Apini
Key to the Species of Apis in Indonesia
1. Distal abscissa of vein M in hind wing present (Fig. 1A) ................. 2
—. Distal abscissa of vein M in hind wing absent (Fig. 1B) ................... 5
2. Forewing hyaline (Fig. 2A); mesoscutellum yellow-brown, rarely
black; drone with tarsi unmodified; worker size moderate, forewing
length 7–9 mm; cavity-nesting species .............................................. 3
—. Forewing fuscous (Fig. 2B); mesoscutellum black; drone with dense
frond-like setae on meso- and metatarsi (Fig. 2C); worker size large,
forewing length 12–15 mm; open-nesting species ...... A. dorsata Fab.
3. Coloration variable but typically clypeus dark grey, antennal scape
dark brown to black, legs dark brown to black, and body setae
ranging from grey to tan or silver; drone cell cap with pore ..............4
—. Clypeus rusty, antennal scape rusty, legs more rusty to reddish
brown, and body setae rather reddish tan; drone cell cap without pore;
Indonesia (Sulawesi) and Philippines (Mindanao) only……………..
…………………………………………………….A. nigrocincta Sm.
4. Coloration not as below; widespread ........................... A. cerana Fab.
—. Coloration largely rubescent, rubiginous (reddish brown) to
rufo-testaceous (reddish yellow), sometimes light testaceous on basal

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 Michael S. Engel: The honey bees of Indonesia (Hymenoptera: Apidae)

leg podites; Malaysia, Brunei, and Indonesia (Sumatra, Java,

Kalimantan) only .............................................. A. koschevnikovi End.
5. Mesoscutellum black; drone with metabasitarsal process; worker size
small, forewing length 6–7 mm; open-nesting species ….................. 6
—. Mesoscutellum light to dark brown; drone without metabasitarsal
process; worker size moderate, forewing length 7.5–10 mm; cavity-
nesting species .............................................................. A. mellifera L.
6. Metatibia and dorsolateral margin of metabasitarsus with black setae;
metasomal terga I and II black to dark reddish brown, infrequently
with reddish brown tints apically on tergum I or basally on tergum II
(Fig. 3A); drone metabasitarsal process short, less than one-half
metabasitarsus length (Fig. 3B) ......................... A. andreniformis Sm.
—. Metatibia and dorsolateral margin of metabasitarsus with white setae;
metasomal terga I and II reddish orange to reddish brown (Fig. 3C);
drone metabasitarsal process long, more than two-thirds
metabasitarsus length (Fig. 3D) (Java only) .................. A. florea Fab.

Figure 1. Hind wings of representative Apis species depicting presence or absence of

distalmost abscissa of vein M (arrow): (A) Hind wing of worker of Apis cerana
Fabricius, (B) Hind wing of worker of A. mellifera Linnaeus. Images not to same scale;
jugal lobes at base of wings folded under (typical of many dried specimens)

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 Treubia 2012, 39: 41-49

Figure 2. Details of Apis s.str. and Megapis: (A) Forewing of worker of Apis mellifera
Linnaeus, (B) Forewing of worker of A. dorsata Fabricius, showing extensive areas of
dark infuscation, (C) Inner surface of drone metabasitarsus of A. dorsata. Images not to
same scale

Figure 3. Differences between species of Micrapis: (A) Dorsal view of metasoma of

worker of Apis andreniformis Smith, (B) Outer surface of drone metabasitarsus of
A. andreniformis, (C) Dorsal view of metasoma of worker of A. florea Fabricius,
(D) Outer surface of drone metabasitarsus of A. florea. Images not to same scale

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 Michael S. Engel: The honey bees of Indonesia (Hymenoptera: Apidae)

Apis (Micrapis) andreniformis Smith. The Black Dwarf Honey Bee is

the native and most common species of Micrapis in Indonesia, ranging
throughout the western half of the country [Greater Sunda Islands
(Sundaland)] (Otis 1996), although putatively only marginally extending
past the Makassar Strait in Sumbawa and westernmost Flores (Hepburn &
Radloff 2011). The species is often found in forests but is in fact
widespread and found in plains as well, but typically not in human
settlements. Like A. florea, nests are constructed in dense thickets and
often shaded.
Apis (Micrapis) florea Fabricius (Not native, introduced). The Red
Dwarf Honey Bee is native to the mainland of Asia, where it is
widespread, extending from the Sudan in Africa all the way to Vietnam
and southeastern China, although natively its western extremes were the
Iranian Plateau and easternmost areas of Arabia. It was historically
introduced to Java and therefore may be found within Indonesia.
Natively, the closest it comes is around the Thai-Malaysian border on the
Malay Peninsula. As with A. andreniformis, nests are often shaded and
built in thickets, although it is not uncommon to find them building nests
in human settlements and from manmade structures. In the wild the
species tends to be found more often in savannah woodlands, forests, or
disturbed areas. As natural habitats are disturbed, A. florea tends to do
fine, while A. andreniformis would be more susceptible to population
declines owing to habitat destruction (much in the same way that A.
koschevnikovi is more susceptible than is A. cerana).
Apis (Megapis) dorsata Fabricius. The Giant Honey Bee is widespread
and common across Indonesia. Nests, like those of the dwarf honey bees,
are exposed. The nests of A. dorsata are typically constructed in open,
conspicuous areas, such as tall trees, rock faces, or buildings, and are
often aggregated and at heights above 6 m, although they can be found
below this level frequently. Workers of A. dorsata can be quite aggressive
when colonies are disturbed and their sting is perhaps the most painful of
any species of Apis. Various authors have argued to split A. dorsata into
multiple species, the most prominent of which is A. d. laboriosa Smith
(e.g., Sakagami et al. 1980, Radloff et al. 2011), but also the forms A. d.
breviligula (Maa) in the Philippines (Maa 1953, Lo et al. 2010) and A. d.
binghami Cockerell in Sulawesi and the Sula Islands (Maa 1953). None
of these have rigorous support as distinct species (Engel 1999).
Apis (Apis) cerana Fabricius. This is, of course, the common Eastern
Honey Bee, which is widespread over most of Asia. Apis cerana, like A.
mellifera, is a robust species which does well in managed and disturbed

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 Treubia 2012, 39: 41-49

systems, in stark contrast to some of its close relatives (e.g., A.

koschevnikovi) which are more susceptible to habitat degradation. It is
also a species with considerable variation, some of which has at times
been split off as separate species (e.g., Maa 1953). Most notably the
subspecies, A. cerana nuluensis Tingek et al. is merely an autapomorphic
type of A. cerana (Engel 1999), showing all of the usual features of a high
elevation form for bees (much in the same manner as distinctive mountain
populations of A. mellifera in Africa). This form is found in the
mountains of Sabah, although it is possible it extends along the entire
Crocker Range and from there deeper into Borneo, perhaps including
Kalimantan.
Apis (Apis) koschevnikovi Enderlein. Apis koschevnikovi is found on the
Malay Peninsula, and throughout Borneo, Sumatra, and Java, although in
many places where the species was once recorded populations have not
been seen recently suggesting that the bees may be in decline or at least
susceptible to human-induced habitat disruption (Otis 1996, Hepburn &
Radloff 2011). The species tends to prefer tropical evergreen forests.
Given the observed declines in populations this species should be studied
carefully and efforts made to preserve the habitats in which it occurs.
Apis (Apis) nigrocincta Smith. Apis nigrocincta occurs on Sulawesi and
Mindanao in the Philippines. Little is known of its biology except that,
where documented, it is similar to that of A. koschevnikovi. Perhaps one
of the most distinctive features of the species is not to be found in the
bees themselves, but instead their nests which lack the well-known pore
in the drone cell cappings, a feature otherwise present in A. cerana
(Hadisoesilo & Otis 1998). This is a little-studied honey bee richly
deserving of intensified research.
Apis (Apis) mellifera Linnaeus (Not native; introduced). The Western
Honey Bee, A. mellifera, has been artificially transported by man for
thousands of years. Its introduction and management in the islands of
Southeast Asia has been largely a phenomenon of more recent efforts, and
mostly by larger commercial interests over those of indigenous and local
peoples. I strongly advocate the position taken by Koeniger et al. (2010)
in advocating for the preservation and development of Asian honeys by
native Asian honey bee species over the introduction of this European
species, and one which has a detrimental effect on the surrounding
environments.

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 Michael S. Engel: The honey bees of Indonesia (Hymenoptera: Apidae)

ACKNOWLEDGMENTS
I am grateful to the numerous beekeepers and honey bee scientists
who encouraged me to prepare this brief report on Apis species, portions
of which are being developed for a world monograph and key to honey
bees, and to Junichi Kojima for a review of the manuscript.
Photomicrographs were prepared with the assistance of
Dr. I.A. Hinojosa-Díaz and with support of the Engel Illustration Fund of
the University of Kansas College of Liberal Arts and Sciences. This is a
contribution of the Division of Entomology, University of Kansas Natural
History Museum.

REFERENCES
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(Hymenoptera: Apidae). Apidologie 29(3): 265–281.
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Apoidea (Hymenoptera). Bulletin of the American Museum of Natural
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Engel, M.S., 2001b. Monophyly and extensive extinction of advanced eusocial
bees: Insights from an unexpected Eocene diversity. Proceedings of the
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Engel, M.S., 2001c. The honey bees of Thailand (Hymenoptera: Apidae).
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Engel, M.S., 2002. The honey bees of India, Hymenoptera: Apidae. Journal of
the Bombay Natural History Society 99(1): 3–7.
Engel, M.S., 2005. Family-group names for bees (Hymenoptera: Apoidea).
American Museum Novitates 3476: 1–33.
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(Hymenoptera: Apidae). American Museum Novitates 3504: 1–12.
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the Centre of Apis Diversity. Natural History Publications (Borneo), Kota
Kinabalu, xix+[i]+262 pp.

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 Treubia 2012, 39: 41-49

Kotthoff, U., T. Wappler & M.S. Engel, 2011. Miocene honey bees from the
Randeck Maar of southwestern Germany (Hymenoptera, Apidae). ZooKeys
96: 11–37.
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(Hymenoptera: Apidae; Apis) in Asia. Journal of the Kansas
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Received: November 15, 2011

Accepted: May 14, 2012

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MacKinnon, J. & K. Phillips, 1993. Field Guide to the Birds of Borneo, Sumatra,
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Maddison, D.R., 1995. Hemiptera. True bugs, cicadas, leafhoppers, aphids, etc..
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 CONTENT
VOL 39, DECEMBER 2012

CONTENT PAGES

Anang Setiawan Achmadi, Ibnu Maryanto and

Maharadatunkamsi. Systematic and descriptions of new species
of Maxomys (Muridae) …………………………….…………..….. 1

Kazuma Matsumoto, Woro A. Noerdjito and Endang Cholik.

Butterflies recently recorded from Lombok ……………………….. 27

Michael S. Engel. The honey bees of Indonesia (Hymenoptera:

Apidae) …………………..………………………………………… 41

Maharadatunkamsi. Morphological variation in

Chironax melanocephalus (Chiroptera: Pteropodidae) from
Indonesia and description of new subspecies ………………….…….. 51

Djunijanti Peggie. A list of the butterflies of Ujung Kulon

National Park, Java, Indonesia …………………….……………..... 67
Awit Suwito, Takahide A. Ishida, Kouhei Hattori and
Masahito T. Kimura. Territorial and mating behaviours of two
flower-breeding Drosophila species, D. elegans and D. gunungcola
(Diptera: Drosophilidae) at Cibodas, West Java,
Indonesia ………………………………………………..…………. 77