2.

BEE SPECIES DESCRIPTION

Bees kept by beekeepers are essentially wild animals and are not domesticated in the way of other
livestock species. In some areas, for example, Europe and Africa, the bees used in beekeeping are
indigenous species, and beekeepers are helping to maintain biodiversity by keeping healthy stocks of
these bees. Until recently, it was true to say that any honeybees kept inside a hive by a beekeeper
would be able to survive just as well living on their own in the wild. However, in recent years, man has
spread honeybee pests and predators around the world, and this means that in some regions, the
indigenous populations of honeybees have been killed and the only bees now surviving are those
managed by beekeepers. For example, in Europe, honeybee colonies can only survive when beekeepers
control levels of the (introduced from Asia) parasitic mite Varroa destructor.

Honey hunting, the plundering of wild nests of honeybees to obtain crops of honey and beeswax, is
practised throughout the world, wherever wild nesting honeybee colonies are still abundant. However, for
thousands of years it has been known that obtaining honey is made much easier and more convenient if
bees are encouraged to nest inside a hive. Apiculture covers this whole, broad range of activities from the
total plundering of wild bee nests for harvests of honey and beeswax, through to ‘conventional’
beekeeping, i.e. the keeping and management of a colony of bees inside a human-made beehive.

BEE SPECIES
In 1988, a bee preserved in amber from New Jersey was identified by US entomologists (Michener and
Grimaldi, 1988). It was a worker, stingless bee of the species Trigona prisca, identical to bees of this
species today. The amber dates from 80 million years ago, and we therefore know that bees of today
were already evolved at that time. There are maybe around 30,000 bee species: about half have so far
been recorded by entomologists. Most bees are solitary, which means that each female bee makes her
own nest, lays a single egg and provides food for the single larva that develops. A few species show a
high level of social development and live together in a permanent, large colony, headed by a single egg-
laying queen. Although many species of bees collect nectar that they convert to honey and store as a
food source, it is only these large colonies formed by social species that store appreciable quantities of
honey. Only a very few species – maybe 30 or so – are exploited by humans for honey production.

These are the honeybees and stingless bees that have been, or are still, exploited by man to varying
extents for their honey stores. Man has exploited them for thousands of years: until recent centuries,
honey was the most common sweetening commodity. There are also a few, very rare instances of
bumblebees being plundered for honey. Of course, the rest of the 30 000 bee species are also plant
pollinators that are vital for the maintenance of biodiversity, and a few of these species are managed
commercially for this purpose.

BEE TAXONOMY
The following is the current view of bee taxonomy according to Michener (2000): all bee species are
classified within seven main families, and one of these is the family Apidae. Apidae has three
subfamilies: Xylocopinae, Nomadinae and Apinae. The subfamily Apinae has nineteen tribes including
Apini (honeybees), Meliponini (includes stingless bees), and Bombini (includes bumblebees). The tribe
Meliponini are the stingless bees found in tropical and southern subtropical areas throughout the
world (see Chapter 6).

The tribe Apini contains just one genus, Apis and these are the true honeybees. Like the Meliponini,
they are social bees that establish permanent colonies. It is these bees’ social behaviour, storing
significant quantities of honey for the colony to survive dearth periods, which means they have been,
and are still today exploited by human societies for their honey stores.

5
 Bees and their role in forest livelihoods

HONEYBEES
There are very few species of honeybees. Most beekeeping textbooks still declare that there are just
four species: Apis mellifera, Apis cerana, Apis florea and Apis dorsata (Ruttner, 1988). The honeybee is
one of the most studied of all animals, other than man, yet this research has been almost entirely on the
European honeybee Apis mellifera. Amazingly however, only within the past 15 years or so a number
of ‘new’ honeybee species have been recorded for science, and Michener names eleven species in the
genus Apis. They are:

Apis andreniformis Apis koschevnikovi

Apis binghami Apis laboriosa
Apis breviligula Apis mellifera
Apis cerana Apis nigrocincta
Apis dorsata Apis nuluensis
Apis florea

These eleven species of honeybees nest in one of two different ways, and this nesting behaviour determines
whether or not the bees will tolerate being kept inside a man-made hive. Some of the species make nests
consisting of a series of parallel combs, other species nest on just one, single comb. The species that build a
series of parallel combs usually nest inside cavities, and this behaviour enables them to nest inside man-made
containers and therefore opens up possibilities for the keeping and management of these bees inside hives.

TABLE 2
Species of honeybees: type of nest
Honeybee species whose nests consist of multiple combs Honeybee species whose nests are single combs
(cavity nesting honeybees)

Apis cerana Apis andreniformis

Apis koschevnikovi Apis binghami
Apis mellifera Apis breviligula
Apis nigrocincta Apis dorsata
Apis nuluensis Apis florea
Apis laboriosa

The species that build single combs usually nest in the open. They cannot be kept in hives and the
single comb behaviour does not lend itself to beekeeping management practices, although the honey
and other products of these species are harvested by some societies.

Honeybee species whose nests consist of multiple combs

Apis mellifera
Other names for Apis mellifera are the hive bee, the European bee, the Western hive bee, and the occidental
honeybee. Most standard beekeeping texts relate only to Apis mellifera (although this is not always stated).

Apis mellifera is indigenous to Africa, Europe and the Middle East. It has been introduced to the
Americas, Australasia and much of the rest of the world. Today, Argentina, China and Mexico have
the largest honey industries in the world, and all are based on the introduced Apis mellifera honeybee.

There are many different races of Apis mellifera, some tropical, others temperate. The Africanised
honeybees in South and Central America are descended from tropical African Apis mellifera. Different
races of Apis mellifera have different sizes of individual bees and colonies. Generally, Apis mellifera are
regarded as the medium-sized honeybees, against which other species are judged as "large" or "small".

Apis mellifera usually builds its nest inside an enclosed space. The nest consists of a series of parallel
combs, and there are typically 30 000-100 000 honeybees in one colony.

6
 Bees and their role in forest livelihoods

Apis cerana
Another name used for Apis cerana is the Asian hive bee, and it is sometimes incorrectly named Apis
indica. Apis cerana is indigenous to Asia between Afghanistan and Japan, and occur from Russia and
China in the north to southern Indonesia. Apis cerana has been introduced recently to Papua New
Guinea. Apis cerana builds a nest consisting of a series of parallel combs, similar in style to Apis
mellifera, and builds its nest within a cavity. As with Apis mellifera, Apis cerana occurs over a huge
geographical area, and it varies in size throughout its range: tropical races are smaller, with smaller
colonies. There are many different races of Apis cerana, as could be expected from the wide range of
habitats it occupies from temperate mountain regions to tropical islands.

Apis koschevnikovi
This honeybee species has been identified only in Sabah, Malaysia in Northern Borneo. Locally known
as the red bee, this species was named for a short period Apis vechti. The individual bees are slightly
larger than Apis cerana found in the same locality, but otherwise the nests of these bees are similar in
size and construction. They are known locally as red bees due to their reddish hue when clustering.

Apis nigrocincta and Apis nuluensis

Apis nigrocincta has been identified only in Sulawesi in Indonesia (Otis, 1996), and Apis nuluensis only
in Borneo. Their nesting behaviour is similar to Apis cerana and Apis koschevnikovi, described above.

Honeybee species whose nests are single combs

Apis andreniformis and Apis florea

These are very small-sized species of bees, and their single comb nests are small too: often no larger
than 150-200 cm wide. Other names include the little honeybee, and sometimes (wrongly) the dwarf
honeybee. These bee species build a single-comb nest, usually fairly low down in bushes, or in the
open, suspended from a branch or (for Apis florea) rock surface. Apis andreniformis has been identified
in South East Asia, Borneo, the Philippines and the southern Chinese peninsula, while Apis florea is
indigenous from Oman spreading southeast through Asia as far as some of the islands of Indonesia and
the Philippines. In 1985, it was identified in Sudan and lately reported in Iraq. However, it is only
recently that Apis andreniformis has been recognised, and some records for Apis florea may prove to be
for Apis andreniformis.

Apis dorsata
Other names for Apis dorsata are the rock bee, the giant honeybee, or the cliff bee. On the western
edge of its distribution, Apis dorsata is found only as far as Afghanistan but its southeast occurrence
extends east of Bali. Its northern distribution is limited by the Himalayas. There is morphometric and
genetic evidence for many different subspecies of Apis dorsata that may eventually be proved separate
species. Apis dorsata bees are large, and their nests consist of single large combs suspended from a
branch, cliff face or building.

Apis binghami and Apis breviligula

Apis binghami occurs in Sulawesi in Indonesia, and Apis breviligula occurs in the Philippines. Maa
(1953) first recorded them as separate species, although subsequent authors ignored this and regarded
them all as the same species, Apis dorsata. Recently, with genetic analysis allowing increasing
understanding of the great diversity with the species Apis dorsata, these two are once again regarded as
separate species.

Apis laboriosa
Apis laboriosa are the largest of the honeybees. They are found in the Himalayas (Nepal, Bhutan, and
China) at higher altitudes than Apis dorsata. Apis laboriosa nests are similar to those of Apis dorsata,
but Apis laboriosa colonies are usually found together in clusters, with sometimes up to 100 combs
suspended from a cliff face very near to one another, although Apis dorsata may also be found nesting
in this way.
7
 Bees and their role in forest livelihoods

BEE SPECIES USED FOR APICULTURE

The honeybees most widely used for beekeeping are European races of Apis mellifera, the species of
honeybee also indigenous to Africa and the Middle East. No species of honeybee occurs naturally in the
Americas, Australia, New Zealand or Pacific islands: European bees have been introduced to these
regions during the last four centuries. Over the last 30 years, European bees have been also introduced
to most countries of Asia. In industrialized countries, all beekeeping technology has been developed
for use with European honeybees, and most beekeeping and research literature relate only to this bee.

Other honeybee species are also exploited by humans for their honey. Although the cavity nesting
species can be kept in hives, and managed according to beekeeping practices, in some countries, wild
nesting colonies of these bees are still sought by honey hunters.

The single-comb nesting species cannot be kept inside hives, so it is only wild-nesting colonies that are
exploited by honey hunting. There are of course exceptions: Apis florea is managed by beekeepers in
Oman (Dutton, 1982), and in several countries in Asia, Apis dorsata is managed to some extent, for
example in India (Mahindre, 2004) and Vietnam (Mulder et al, 2001). There is more information on
this in Chapter 5.

DIFFERENCES BETWEEN TROPICAL AND TEMPERATE ZONE RACES OF HONEYBEES

European races of Apis mellifera have evolved in temperate climates with long, cold winters when little
or nothing is in flower. They store honey to serve as a food supply to survive these times of dearth
when there is little or no food available. Apart from swarming (the colony’s reproduction), they
remain in their hive because they are unlikely to survive if they leave in search of a new nesting place.
By comparison, all tropical races and species of honeybees are far more likely to abandon their nest or
hive if disturbed, because in the tropics they have a reasonable chance of survival. In some areas,
tropical honeybee colonies migrate seasonally. These are crucial factors making the management of
tropical honeybees different from the management of temperate zone honeybees.

TABLE 3
Species of honeybees: indigenous distribution
Region Indigenous honeybee species Honeybee species introduced
AFRICA Apis mellifera Apis florea introduced to Sudan, 1985
ASIA* Apis andreniformis Apis mellifera
Apis binghami
Apis breviligula
Apis cerana
Apis dorsata
Apis florea
Apis laboriosa
Apis koschevnikovi
Apis nigrocincta
Apis nuluensis
AUSTRALASIA No indigenous honeybees Apis mellifera
Apis cerana has been introduced to Papua
New Guinea
EUROPE Apis mellifera
MIDDLE EAST Apis mellifera
Apis florea
THE AMERICAS No indigenous honeybees Apis mellifera
* Not all of these species are indigenous to every country of Asia.

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 Bees and their role in forest livelihoods

AFRICA
Apis mellifera honeybees are indigenous to Africa. There are many different races of African bees; see
Ruttner (1998) for more information. In South Africa bees are of the race Apis mellifera capensis, a race
of bee with unique biology and behaviour (see below). Tropical races of Apis mellifera are slightly
smaller than the European races of Apis mellifera and they have different biology and behaviour: they
are readily alerted to fly off the comb and to defend themselves. In many African countries, local
beekeeping methods are used, with log, bark, basket or clay hives placed in trees. Where the behaviour
of bees is to swarm and migrate, it can be a good beekeeping strategy to use a large number of low cost
hives. This means that the beekeeper can afford to have a large number of hives and accept that some of
them will be unoccupied at some periods. Throughout Africa honey hunting from wild nests is carried
out wherever sufficient natural resources remain. Stingless bees are also present throughout tropical
and southern sub-tropical Africa.

BOX 3
Apis mellifera capensis
Apis mellifera capensis, known as the Cape honeybee, is a race of Apis mellifera whose natural distribution is confined
to the southern tip of Africa, and which has a unique, highly complex biology that has only recently been understood.
The unique feature of Apis mellifera capensis is that worker bees, without any mating taking place, are able to lay
diploid, female eggs. This biology is not known in any other honeybee species or race, where the usual ‘rule’ is that
worker bees lay only haploid, male eggs that develop into drones.

The recent (1990) movement by beekeepers of these bees from southern to northern South Africa caused the
widespread death of African honeybee (Apis mellifera scutellata) colonies. The Apis mellifera capensis workers enter
the Apis mellifera scutellata colonies, and this soon leads to colony break down and death. It seems that the eggs laid
by the Apis mellifera capensis bees evade being killed by other worker bees, as would normally happen, and ultimately
the colony breaks down. The spread of these Apis mellifera capensis bees in South Africa, together with the recent
introduction of Varroa mites, has severely curtailed beekeeping in South Africa and these issues may eventually affect
on bees and beekeeping throughout Africa.

ASIA
At least eight honeybee species, varying in biology and behaviour, occur naturally within Asia. Some
of these bee species build nests consisting of single combs, in trees, bushes, or in cliffs, and a great
variety of methods have been developed by human societies for their exploitation.

For example, the giant honeybee, Apis dorsata, suspends its large combs (often one metre in diameter)
from tree branches and overhanging ledges on rocks and buildings. Man obtains honey crops from this
species by plundering their colonies, and this activity is known as honey hunting. Throughout Asia,
from Gurung tribesmen in the Himalayas, to mangrove-dwellers in the Sunderbans of Bangladesh, the
rain-forest people in Malaysia, people living in the river deltas of southern Vietnam, and indeed,
wherever the giant honeybee is present, honey hunters have their own customs for exploiting these
bees (see Chapter 5).

Apis cerana is known as the Asian hive bee because like European Apis mellifera, it can be kept and
managed inside a hive. Moveable frame hives and movable comb hives (top-bar hives) have therefore
been developed for Apis cerana and the other cavity nesting hive bees.

Stingless bees are also present throughout tropical and southern sub-tropical Asia.

European Apis mellifera have been introduced to most of Asia as shown in Table 4, and this exotic
species may now be the predominant honeybee species present in China, Japan and Thailand, and
other countries of Asia.

9
 Bees and their role in forest livelihoods

TABLE 4
Numbers of Apis mellifera colonies in Asia
1984 1994 2004 1984 1994 2004

Afghanistan 20 000 ? Japan 284 000 225 000

Bangladesh 0 ? Malaysia <500 present

Bhutan 0 50 Nepal 2 1 000+

Brunei ? 0 Pakistan 1 000 14 000

Burma 2 000 2 000+ 5 000 Philippines 2 000 6 000

Cambodia ? ? Singapore ? present

China 4 000 000 6 800 000 South 280 000 300 000 790 000
Korea
Hong Kong ? 100+ Sri Lanka 4 not
permitted
India 3 000 80 000 Thailand 30 000 100 000 300 000

Indonesia 1 000 31 000 Vietnam 16 000 70 000 470 000

Laos ? present

AUSTRALASIA AND PACIFIC OCEAN ISLANDS

There are no honeybees indigenous to this region, although there are indigenous species of stingless
bees that have been harvested traditionally. European races of Apis mellifera have been widely
introduced and are used for beekeeping. Recently Apis cerana has been introduced to Papua New
Guinea.

CARIBBEAN
Although indigenous stingless bees are present, no honeybees are naturally occurring in these islands.
Apis mellifera of European origin have been introduced to most of them and beekeeping industries
have developed using European-style beekeeping methods. With the rapid spread of honeybee diseases
around the world, it is increasingly important that these islands endeavour to maintain stocks of
disease-free bees. Caribbean beekeepers must watch for Africanised bees that have already arrived in
Trinidad.

EUROPE
Apis mellifera is the honeybee indigenous to Europe, and there are many different races of the bees. See
Ruttner (1988) for a detailed account. During the 20th century, bees were moved by beekeepers from
one area to another and many hybrids were created. Today there is more interest to identify and
preserve the original races of bees that are now appreciated to be the bees best suited for their own
areas. For example, Slovenia is home to the indigenous Carniolan bee Apis mellifera carnica, known as
“sivka” meaning “grizzly” because of the bright grey hair along the edges of its abdomen, and admired
by beekeepers for its characteristic gentleness and diligence. Because of this behaviour, people started
to keep it in hives close to home. News of the gentle character of this grey bee soon spread to other
nations and by the end of the 19th century; there was the beginning of a lively trade in live bees and
swarms, later to include Carniolan queens. Until the beginning of World War I, specialized Slovene
merchants exported tens of thousands of bee colonies and, in many places; these completely replaced
the indigenous dark bee. Today, honeybee queen breeders, who sell approximately 40 000 queens,
mostly to the countries of Central and Western Europe, with exports increasing annually, are
continuing their work. Slovenia joined the EU in May 2004, and the beekeeping sector was well
prepared, with legislation for an “Authentic Carniolan Trademark” for the marketing of indigenous
Carniolan genetic material and a well-organised reserve area for the indigenous bees.

10
 Bees and their role in forest livelihoods

Apis mellifera carnica is also kept fruitfully in neighbouring Austria and Croatia, as well as elsewhere
in Central and Eastern Europe. This bee species is well adapted to the climate and foraging conditions
of these countries. It tolerates local conditions: cold, snowy winters, frequent rainy and windy
summers and makes good use of available forage. One of its beneficial characteristics is discovering and
collecting honeydew from spruce and fir trees. Almost 60 percent of Slovenia retains its forest cover,
with mixed coniferous and deciduous forests offering rich forage for bees. The most important honey-
producing trees are fir and spruce, followed by sweet chestnut, lime, sycamore and wild cherry.

BOX 4
Save indigenous bees in Europe1

One of the last remaining populations of the European honeybee Apis mellifera mellifera is threatened. These are the
Black Bees on the Danish Island of Læsø, an isolated island that lies west of Sweden in the Kettegat Sea. In 1992
Denmark signed the Rio Convention on Biological Diversity, and the law was passed for Læsø Island to become a
protected area where only beekeeping with the Black Bees is allowed. After this, beekeepers who kept other bees
claimed compensation, although this claim was later dropped. They also took their case to the European Court in
Luxemburg, but were unsuccessful. The Court ruled that the Preservation Order on the Læsø Black Bee was a
requirement of The Danish Government, and that no other race of bees should be allowed on to the Island. Today on
Læsø there are about 30 beekeepers using the Black Bees, and just a few who continue to fight the ban and illegally
use other bees, and even import bees. This has lead to the recent introduction of Varroa and Acarapis mites.
Ironically, it was only in September 2004 that SICCAM (The International Organization on the preservation of the
Northern European Black Bee) held its biannual conference on Læsø, to focus attention on the need to protect this
special bee population. SICCAM passed a resolution calling for this unique population of bees to receive the protection
it needs.
Now, however, the Danish Minister of Agriculture and Food, Hans Christian Schmidt has decided that it is in the
interests of human liberty for the few, vocal, beekeepers who request it, to be allowed to take in other races of bees to
the Island, and that only a small part of the Island will be a protected area for the Black Bees. The island of Læsø is
only 25 km long; therefore, as every beekeeper will understand, it is not possible to keep the populations of bees
separate.
Meanwhile, the Danish Beekeepers Federation has fought hard to protect the black bees, even though its own
government subsidy is at stake.
The majority of beekeepers in Denmark want the Black Bees on Læsø to be protected. This is a precious resource, not
just for Denmark but also in world terms.

THE AMERICAS
There are no honeybees indigenous to the Americas. Instead, their ecological niche was filled by the
many different species of stingless bees, which were, and still are in some areas, exploited for their
honey that is especially valued for its medicinal properties. Knowing nothing of these indigenous bees,
European settlers long ago took with them European bees, and an industry developed based on this
bee. In 1956, some tropical, African Apis mellifera bees were introduced into Brazil. These bees
survived far more successfully in tropical Brazil than their European Apis mellifera predecessors. These
'Africanised' bees (dubbed 'killer bees' by the media) have spread through tropical parts of South and
Central America, and are now in southern USA. In Brazil and neighbouring countries, beekeepers
developed new management methods and now make excellent livelihoods with these bees.

THE NEAR EAST

Apis mellifera is also the indigenous bee of the Near East, and as everywhere, there are indigenous
races of Apis mellifera that have their own characteristics highly suited to local conditions. Middle
Eastern races include Apis mellifera syriaca and Apis mellifera yemenitica, desert races that survive hot,
arid conditions. Apis florea is also present in some countries of the Middle East, and its honey is highly
prized, often changing hands at over US$100 per kilogram.

PROBLEMS WITH THE INTRODUCTION OF EXOTIC BEE SPECIES AND RACES

As far as beekeepers are concerned, throughout the 20thcentury the other man’s grass was always
greener – bees in other countries were viewed as more prolific, gentler, more disease resistant, less
prone to swarming, more yellow, blacker. Indeed many beekeepers still think this way, and this has led

1
Bradbear, 2005.
11
 Bees and their role in forest livelihoods

to the disasters of recent years, when races of bees, or diseases and parasites of honeybees have been
spread around the world with serious consequences for the beekeeping industries, and indigenous
populations of bees, in many countries. This has been caused entirely by the movement of honeybee
colonies by man.

For example, the mite Varroa destructor is a ‘natural’ parasite of Asian honeybees that survive in the
presence of the mite. However, when particular races of the mite are introduced to European Apis
mellifera honeybees (the bee used for beekeeping in most industrialized countries), the whole colony
will be killed unless action is taken by the beekeeper. These mites have now been introduced to many
beekeeping countries and, for example, most populations of wild honeybees throughout Europe have
been killed during the last 20 years or so. Mites become resistant to medicines developed for their
treatment, and research is underway in many countries to find better, integrated control methods, or
resistant strains of bees.

Recently another predator, the small hive beetle, Aethina tumida, has been spread from Africa (where
it is a relatively harmless pest for bees) to honeybee colonies in the USA, where it leads to destruction
of European honeybee colonies.

The introduction of African bees to south America was initially viewed as a disaster, as the introduced
African bees survived very well in their new habitat, and their population quickly expanded through
south and central America, replacing existing populations of European honeybees, there were less well
suited to the tropical environment. However, today some view this amazing, dramatic event in a more
sympathetic light – as beekeeping industries have learned to adapt to the African bees. The Brazilian
scientist who introduced the African bees, Professor Warwick Kerr, has with hindsight, expressed the
opinion that it would have been wiser to have focussed efforts on the Americas’ indigenous, stingless
bees (Bradbear, 1993).

Honeybees and used beekeeping equipment must never be moved from one area to another without
expert consideration of the consequences. Just a very few regions remain without introduced honeybee
diseases, and these are mainly in developing countries. It will be highly beneficial for these countries if
they can retain their stocks of disease-free honeybees: they may in the future be able to market their
disease free stocks, or export disease free queen bees, and it makes possibilities for organic honey and
beeswax production cheaper and easier.

THE CONSERVATION OF INDIGENOUS HONEYBEE SPECIES AND RACES

Globalisation is taking place in beekeeping, as in every other sector. Beekeeping with European races
of honeybees, plus all associated technology, is being spread around the world. The consequences of
competition between introduced (exotic) honeybees and indigenous honeybee species and races are
unknown.

12
3. THE IMPORTANCE OF BEES IN NATURE
BEES AS PART OF ECOSYSTEMS
Pollinators strongly influence ecological relationships, ecosystem conservation and stability, genetic
variation in the plant community, floral diversity, specialization and evolution. Bees play an important,
but little recognized role in most terrestrial ecosystems where there is green vegetation cover for at
least 3 to 4 months each year. In tropical forests, savannah woodlands, mangrove, and in temperate
deciduous forests, many species of plants and animals would not survive if bees were missing. This is
because the production of seeds, nuts, berries and fruits are highly dependent on insect pollination, and
among the pollinating insects, bees are the major pollinators. In rain forests, especially in high
mountain forests where it is too cold for most bees, other pollinators like bats and birds play a greater
role in plant pollination. In farmed areas, bees are needed for the pollination of many cultivated crops
(see Chapter 7), and for maintaining biodiversity in ‘islands’ of non-cultivated areas. The main role of
bees in the different ecosystems is their pollination work. Other animal species are connected with
bees: either because they eat the brood or honey, pollen or wax, because they are parasitic to the bees,
or simply because they live within the bees nest.

WHAT IS POLLINATION?
Pollination is transfer of pollen from the anther (the male part of the flower) to the stigma (the female
part of the flower). Some plants can pollinate themselves: in this case, the pollen passes from the anther
to the stigma inside the same flower, and this is called self-pollination. Other plants need pollen to be
transferred between different flowers or different individuals of the plant. This is cross-pollination.
Many plants can be pollinated both ways. Plants can be pollinated by wind or animals.

Some plants have only one method for pollination, others use a combination. The knowledge of
pollination by animal pollination (Zoophily) in the tropics is still little known, and much work and
research have to be done in this area. Some general rules can be used to detect whether a plant is
pollinated by bees, flies, beetles, wasps, butterflies, moths, thrips, birds, bats, marsupials, slugs or
rodents. Flowers pollinated by bees most often bloom in daytime, they can have different colours, but
seldom red. The scent of daytime bee pollinated flowers tends to be less strong than that of night-
pollinated flowers, often pollinated by bats or moths. Honeybee pollinated flowers have nectar tubes
not more than 2 cm long. They have nectar guides (patterns to direct the bee towards the nectary) and
often a landing place for bees. Bees are especially attracted to white, blue and yellow flowers. Plants
pollinated by insects are called “entomophilous”, and insects are generally the most important
pollinators.

THE POLLINATION WORK OF BEES

If we look at the many colourful and different looking flowers, we should not forget that they have
developed as an adaptation for the bees and other pollinators, and not to please humans! Bees and most
flowering plants have developed a complex interdependence during millions of years. An estimated
80 percent of flowering plants are entomophilous i.e. depending more or less on insect pollination to
be able to reproduce, and it is estimated that half of the pollinators of tropical plants are bees.

The efficiency of honeybees is due to their great numbers, their physique and their behaviour of
foraging on only one plant species at one time. The bees have to find their food in flowers. The food
can be nectar or pollen. Nectar is produced to attract the bees. Pollen is also attracting the bees, but it
has another function too: it is produced to ensure the next generation of plants. Bee pollinated flowers
have evolved in such a way that a visiting bee has to brush against the flower’s anthers bearing pollen,
or there may be a special mechanism to release the anthers to spring up or down to cover the bee with
pollen. Compared with other insects, bees are extremely hairy. Each hair has a branched structure that
makes it highly effective at catching pollen.

13
 Bees and their role in forest livelihoods

While flying to the next flower, the honeybee will brush herself and move many of the pollen grains, to
arrange them in the pollen baskets made of stiff hairs on her hind legs. Some pollen grains are so dry that they
cannot be formed into a clump. To prevent the pollen falling off during flight, the bee will regurgitate some
nectar and mix it with the pollen. This gives the sweet taste when eating pollen balls collected by bees. It also
makes the pollen a little darker so that it can be difficult to see from which plants it comes. Some bees do not
have pollen baskets – they transport the pollen in the hair on their abdomen (e.g. Osmia bees and leaf cutter
bees). When the honeybee with pollen is landing in the next flower, there will be pollen enough left on the
bees’ body hairs to pollinate the new flower, by delivering some grains to the flower’s stigma. Now
pollination has taken place. To create a seed, the pollen grain has to grow a small tube inside the stigma to the
ovary of the flower. Then a male gamete can travel through the tube, fertilize the egg cell and start
development of the fertile seed. Now the fertilization has taken place.

Some plants need several successful visits from bees to ensure that all the flower’s eggs are fertilized.
For example, some varieties of strawberry need about 20 pollen grains – requiring visits by several
bees, an apple flower may need four or five bee visits to receive enough pollen grains for complete
fertilisation. If the fertilization is inadequate because of lack of bees, not all seeds will develop, and the
shape of the fruit will be poor and small. Fertilization is the beginning of a new seed, which perhaps
will grow and develop into a new plant. The new plant will bloom, provide the bees with food, be
pollinated, and be fertilized, and in this way, the story continues.

The forager bee returns to the honeybee colony with her pollen loads, which are placed in the nest in
areas of comb close to the brood.

Bees have to learn where in a flower the nectar is to be found. To guide the bees, many plants have bee-tracks,
which are lines of colour leading the bee towards the nectar. These can sometimes be seen by humans, but
some are in the ultra-violet part of the spectrum and visible to bees, but not humans. In this way, the plant
also guides the visiting bee to pass the anthers or stigma in the right way. Bees have no problems in finding the
nectar in flat, open flowers, but in flowers that are more complex, they have to learn it by trial and error. After
some visits in the same type of flower, the bee has learned where the nectar is, and learns this for the next visit.
Pollen is the protein food for bees. Without pollen, the young nurse bees cannot produce bee milk or royal
jelly to feed the queen and brood. If no pollen is available to the colony, egg laying by the queen will stop.

Usually a honeybee can visit between 50-1000 flowers in one trip, which takes between 30 minutes to
four hours. In Europe, a bee can make between seven and 14 trips a day. A colony with 25,000 forager
bees, each making 10 trips a day, is able to pollinate 250 million flowers.

The ability of the honeybee to communicate to other bees in the colony where to go for collecting more
pollen and nectar is very important for their efficiency as pollinators. When a scout bee has found a good
nectar or pollen source, she will return to the colony and communicate to other bees where they can find
the same food. This is done with a special dance indicating the distance, quality, and direction from the
nest. Flowers closer than around 200 metres are just announced with the waggle dance without indicating
any direction. Chapter 6 describes how these stingless bees are guided to the flowers.

When bees begin foraging for pollen and/or nectar, they will visit the same species of flowers and work
there as long as plenty of nectar or pollen can be found. For example, if a honeybee starts collecting in
an Acacia tree, she will fly from Acacia flower to Acacia flower, and not behave as many other insects
do, visiting different species of plants within the same trip without any great pollination effect. This
behaviour of bees is called foraging constancy.

Some flowers are open and with nectar all day and night, but others are open only for a few hours in
the morning, afternoon or night. The single worker bee learns and remembers what time the different
flowers are worth visiting. One bee can remember the opening time for up to seven different types of
flowers. The honeybees are pollinating a great number of different plant species, and they do it
effectively. Some solitary bee species are much more specialized for pollinating specific plant species.
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 Bees and their role in forest livelihoods

SPECIALIZED POLLINATION
Some species of plants and bees have developed a close interdependence in connection with
pollination. Such a mutual adaptation and interdependence between a plant and pollinator is a result of
a long and intimate co-evolutionary relationship. The pollinating bees of the Brazil nut tree
Bertholletia excelsa is an illustrative example of such a relationship and its economic importance.

The Brazil nut tree grows wild in the Amazon Forest. Brazil nuts are one of the economically most
important wild products growing trees in the area, with more than 50 000 tonnes of the nuts exported
from Brazil every year. The Brazil nut trees cannot be grown in plantations, because they need to be
pollinated by one special bee species, the small shining Euglossa bee. This bee is dependent on the
presence of an orchid species that is found only in the rain forest. They are also the only pollinators for
a number of orchids in the forest. In some species of Euglossa, the male bee collects some scented
material from the flower, which they distribute to attract other males – who do the same and multiply
the effect with a scented cloud, in the end so strong, that it attracts female bees so that mating can take
place. During the collection of the scented material, male bees transfer pollen from orchid to orchid
and pollination takes place. The female Euglossa bees live from nectar from the Brazil nut tree and
pollinate it. This means that without the orchids, there would be no Euglossa bees and no Brazil nut
trees, and none of the many other plants, insects and animals associated with that tree – including the
people whose livelihoods include collection and sale of the Brazil nuts.

Studies in the Amazon forest have shown that many Euglossa bees do not cross open areas. That means
that great parts of forest lose its pollinators when the forest is cut, and open parcels of land are created
between remaining forest islands.

This example is only one of many important specialized interrelations between bees and trees. In spite
of this, the bees perhaps play a minor role as pollinators in the rain forest compared to their role in
temperate forests, monsoon forests and savannah woodland. In tropical rain forests, many trees are
pollinated by birds, bats and insects other than bees. Animal pollination is of greatest importance,
because there is no wind between the trees and because the distance between trees of the same species
may often be great. In that way, it is most convenient for the trees to use animals as pollination vectors.
In tropical forest, there may be rather few flowering plants on the ground because of the trees’ shade.

In European deciduous forests, the forest floor can be totally covered by flowering plants in
springtime, before the trees produce their leaves. These plants often need fast pollination from a great
number of honeybees. Not many other insects are present in high numbers in early spring.

In Denmark, it is seen by forestry people that the presence of bees in forest areas helps to protect the
newly planted trees from being eaten or spoiled from gnawing by roe deer, compared to other
plantations with no bees. The reason is because bees secure a better pollination and seed production of
so many other plants, which the roe dear can forage on instead of the tree seedlings. By pollinating
trees, bushes and herbaceous plants, the bees are important for the food production of all the other
animals and birds in the forest ecosystem dependent on it for food berries, seeds and fruits.

BEES ARE GOOD FOR TREES AND TREES ARE GOOD FOR BEES
Bees and trees belong together. The honeybees and stingless bees have originally developed in forest
biotopes. Given the choice, wild honeybees chose nesting places in trees rather than in an open
landscape. Most often the honeybees prefer to build their combs or nests high in trees instead of close
to the ground, but bees nests can be found everywhere in a tree. In savannah areas with bushfires in the
dry season, a high nesting place is an advantage. When beekeeping is present in a forest, the beekeepers
will be interested in protection of the forests and especially the tall trees preferred by the bees. When
enough bees are present in a forest, they provide a better pollination that leads to improved
regeneration of trees and conservation of the forest’s biodiversity.

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 Bees and their role in forest livelihoods

BEES AND BIODIVERSITY

Without bees there would be no flowering plants, and without flowering plants there would be no
bees. Without bees biodiversity would not be so great. Biodiversity is measured as the number of
different plant and animal species found in a certain unit area. Biodiversity is highest in tropical forest
areas and lowest in the Arctic. High biodiversity is related to the high age of the ecosystem, and a
stable environment. A stable environment creates the possibility of development of specialization and
use of narrow ecological niches. The explanation of the high biodiversity in tropical forests can be as
the species’ efforts to avoid attack by diseases and pests. Both can be much more serious in a tropical
forest biome with a constant supply of water, and a hot and stable temperature. The high diversity
with its high specialization in pollination relationships can also be a danger for the forest. The specialist
pollinator must have access to food all year round. Many of the smaller trees flower all year round or
nearly all year, but the larger trees have blooming seasons. Some flower every year, others every third
or fifth year, where all trees from the same species bloom at the same period and maybe even at the
same hours. If the specialized bees loose their stable resources by tree cutting, they will not be there
when the bigger trees require their pollination service.

The reproduction of plants is simplest as vegetative reproduction – a new tree could just come from a
root shoot. The new tree would then be genetically identical with the mother tree. Vegetative
reproduction alone would be no problem if the environment were stable, but most environments are
not stable over time, they change. It can be climatic changes, new diseases or pests. To be able to adapt
to environmental changes there need to be genetically different plants. In that way there will always be
some plants, which are better adapted than others because of special genetic constitutions. The only
way to constantly mix the genes for the plants is by cross-pollination, where pollen from one plant is
transported by bees to another so that the offspring become genetically different. In that way, there is a
greater chance for at least some of the offspring to survive in the competition of life. In this we find the
bees as one of the most important factors.

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