Chapter 2
Historical Perspective
2.1 The Origins of Honeybees
Honeybees have evolved from short-tongued, spheciform wasps and first appeared
during the Cretaceous period about 130 million years ago. At that time, present-day
continents such as Africa, India, South America, Australia and Antarctica constituted
a single landmass called Gondwana. Germinating in the warm dry Gondwanan climate, flowering plants called angiosperms developed colours and petal patterns to
attract insects, which were more reliable than wind to transfer pollen. In addition to
pollen, flowers eventually produced nectar, providing carbohydrates to their winged
vectors. About 120 million years ago, the honeybee developed its morphologies
specifically to collect pollen and nectar, such as increased fuzziness, pollen baskets,
longer tongues and colonies to store supplies.
As Gondwana gradually broke apart and temperatures cooled dramatically during the Oligocene–Miocene about 35–40 million years ago, European honeybees
went extinct, while Indo-European honeybees survived and began to speciate. Opennesting honeybees perhaps evolved before cavity-nesting bees, probably in India,
but evidence is still lacking. In any event, a cavity-nesting honeybee spread east
and north about 6 million years ago. During a Pleistocene warming about 2–3 million years ago, this bee spread west into Europe and then into Africa to become A.
mellifera.
It is thought that Apis florea and Apis dorsata may have existed as separate species
as early as the Oligocene period. It has not been possible to estimate when bees of
the mellifera/cerana type first appeared on Earth. Mellifera and cerana must have
acquired separate identities during the latter part of the Tertiary era. The two species
were apparently physically separated at the time of the last glaciation, and there was
no subsequent contact between them until that brought about by human intervention
in recent times. In the post-glacial period, mellifera and cerana have shown similar
evolution into geographical subspecies or races.
D. P. Abrol, Asiatic Honeybee Apis cerana,
DOI 10.1007/978-94-007-6928-1_2, © Springer Science+Business Media B.V. 2014
39
40
2 Historical Perspective
2.2 The Development of Subspecies
Although it has long been known that there are many kinds of honeybees, and these
have been the subject of scientific study for more than two centuries, only in recent
years has a comprehensive classification been attempted, which takes into account
not only differences in physical characters between subspecies and their present
geographical distribution but also the geological evidence pointing to their origins
and to the course of their subsequent evolution and distribution.
Like the stingless bees, honeybees first evolved in tropical conditions. The fossil
record shows that at the time the area of land that is now Europe had a tropical climate.
As the climate became cooler, the open-nesting types would not have been able to
survive except by migrating to the tropical region of Southern Asia. For the greater
part of the Tertiary era, Africa was isolated from Europe by sea, and no Tertiary types
of honeybee reached Africa even after a land bridge was established. It is likely that
the development of advanced thermal homeostasis in honeybees, which permitted the
occupation of cool temperate zones, therefore occurred in Southern Asia, possibly
in the Himalayan region. Once established, the cavity-nesting cerana–mellifera type
would spread East and West, eventually occupying both tropic and cool temperate
zones.
A physical separation into two groups probably took place as a result of the
glaciations, which occurred during the Pleistocene period (1 million to 25,000 years
ago) and desert and semi-desert then kept the two groups separate during intervening
warm periods. Thus, mellifera and cerana, although originating from a common
stock, evolved into distinct species. The ultimate Western boundary of the cerana
territory was in Afghanistan some 600 km to the East of the nearest mellifera colonies
in Iran. The cerana territory comprised the Indian Subcontinent south of the great
mountain ranges, Ceylon, Malaysia and Indo-China, and the East Indies including
the Celebes, Timor and the Philippines. In Eastern Asia, it reached a latitude of
46 degrees, and occupied Japan except for the island of Hokkaido. Mellifera spread
westwards through Asia Minor to colonise the Balkans and the Mediterranean region
and southwards through the Arabian Peninsula to occupy central and southern Africa.
Similarities between neighbouring subspecies suggest that the Iberian Peninsula and
southern France were colonised from North Africa.
How far mellifera bees may have penetrated into northern and western Europe
during the warm intervals between the glaciations of the Pleistocene period can only
be a matter of conjecture; what is certain is that no honeybees could have existed
north of the Mediterranean region, the Iberian Peninsula and southwestern France
at the time of the most recent Ice Age. Although at its maximum extent in western
Europe some 18,000 years ago, the ice sheet only reached as far as northern Britain,
the area for hundreds of miles to the south was inhospitable tundra.
In the warm period, which followed the Ice Age (starting about 14,000 years ago),
the ice sheet gradually retreated and the tundra was replaced by forests of birch, pine,
hazel, elm and broad-leaved oak. The western honeybee was once more able to extend
its domain in Europe. In the east, advance beyond the Caucasian region proved
2.2 The Development of Subspecies
41
impossible, owing to the lack of suitable nesting sites in the steppes of southern
Russia. The bees of the Balkan area spread northwards to occupy the eastern Alpine
valleys, central Europe as far as the 50th parallel of latitude, and the western shores
of the Black Sea. In the West, the bees, which had found refuge in southern France
during the Ice Age, spread across Europe north of the Alps, eventually occupying an
area from the Atlantic seaboard to the Ural Mountains. The northernmost limit of
the territory may have been in southern Norway; honeybee remains dating from c.
1200 have been found in an archaeological dig in Oslo, although honeybees had not
been reported in Norway prior to the nineteenth century. The mountain ranges of the
Alps and the Pyrenees obstructed the northward movement of the bees in the Italian
and Iberian peninsulas.
In colonising this vast territory, stretching from the Urals to the Cape of Good
Hope, A. mellifera had to adapt itself to a large variety of habitats and climates ranging
from the continental climate of eastern Europe with its harsh winters, late springs
and hot, dry summers, through Alpine, cool temperate, maritime, Mediterranean,
semi-desert and tropical environments. This adaptation was achieved by natural
selection, producing some two dozen subspecies or races. All the subspecies of the
mellifera group can interbreed given the right conditions, but the crosses show hybrid
characters.
Although cerana bees must have shared a common ancestor with mellifera, they
have evolved into separate species. It is not possible to cross cerana with mellifera
even using instrumental insemination, because the two species are now genetically
incompatible and viable eggs do not result from the cross-fertilisation Other differences include their differing reactions to diseases, infestations and predators. Cerana
can tolerate varroa and has developed an effective defence strategy against the Giant
Hornet, against which mellifera bees have no defence. Cerana is, however, highly
susceptible to the acarine mite, which arrived with the introduction of mellifera bees
into cerana territory. It is also highly susceptible to sac brood and foulbrood, but not
markedly so to nosema.
The different races of A. mellifera can generally be differentiated in physiological
terms. Bees from warmer climates tend to be smaller in size and lighter in colour than
those adapted to the colder regions, although this rule is not invariable. The effect
of altitude seems to be similar to that of increasing latitude. Accurate differentiation
between races of similar appearance requires precise morphometric examination of
representative samples of bees. There are also differences between races in natural
history and biology. Some subspecies are more prone to swarming than others, some
produce large numbers of young queens when swarming, others only a few. Tropical
honeybees frequently ‘abscond’ or migrate, sometimes because of lack of forage
through drought or other causes, perhaps as a defence against predators. Heavy
predation is also a likely cause of the vigorous defence reaction of some races, for
example, the bees of tropical Africa.
The bees of the warmer regions do not need to cluster as tightly as those confined
to the nest through long, cold winters. Brood rearing is adapted to take maximum advantage of the local flora. Where bees of the same race have occupied different kinds
of habitat, they have formed local strains which have accommodated themselves to
42
2 Historical Perspective
the different conditions. Similarly, honeybees of different races which have occupied similar habitats have evolved similar behavioural characters. Even the ‘dance
language’ by which honeybees communicate information about the location of food
sources may differ in detail between races, as different races may be conditioned
to foraging over different distances from the nest. (Professor Goats described these
differing dance patterns as ‘honeybee dialects’.) The behavioural characters of the
different races and strains, brood rearing pattern, foraging behaviour, clustering, etc.,
are fixed genetically, so that a colony cannot readily adapt itself when transferred to
a different kind of environment.
The dark European honeybee, A. mellifera mellifera, is fairly uniform over its
whole range, having had but a comparatively short time in which regional varieties
could evolve, but even in this race, differences can be observed between strains.
In France, where the bee has been domiciled longest, there are distinct differences
in brood rearing pattern between the mellifera bees of the Landes district in the
southwest, the bees of the Paris area and those of Corsica. The Landes bees are
typical ‘heather bees’, conditioned to a principal nectar flow in late summer and
early autumn. In the Paris area, there is no summer nectar flow and the bees show
early spring brood activity. Exchange of colonies between the Landes and Paris
resulted in poor performance in both cases. In Corsica, the mellifera bees follow a
Mediterranean pattern with little or no brood production in summer and a second
peak in autumn.
The behavioural patterns which have evolved in the different races have ensured
the survival of the various subspecies in their native habitats, and some of these
patterns may be repeated in different races. There is one race which, in spite of small
economic importance, possesses an apparently unique biological character, which
renders it of great importance in the study of the genetics of honeybees. In all other
races, when a colony is rendered queenless, laying workers may appear which are
capable of laying drone eggs only. In A. mellifera capensis, the Cape bee, when a
colony is deprived of its queen, a laying worker appears within a few days which,
for a period, is able to lay predominantly diploid worker eggs. From these eggs, true
queens capable of being mated can be raised, re-establishing queenrightness in the
colony.
The genus Apis can be divided into three branches based on their nesting pattern:
the open-nesting single-combed giant honeybees A. dorsata and Apis laboriosa; the
dwarf honeybees A. florea and Apis andreniformis; and the cavity-nesting A. cerana,
Apis koschevnikovi, Apis nuluensis, Apis nigrocincta and A. mellifera. All the nine
species thrive well in environmental extremes like deserts, rain forests and tundra,
but most people are familiar with A. mellifera only, which is known as important
pollinator for agricultural production all over the world.
Early civilizations quickly mastered honey hunting skills, shown in rock art in
Africa, India and Spain. Egypt, Greece, Italy and Israel developed organized beekeeping centres until the Roman Empire dissolved in approximately 400 ad. Christianity
monasteries and convents then served as apiculture centres until Henry VIII closed
them at the beginning of the Reformation. Science and technology provided the next
insights into apiculture during the Enlightenment.
2.3 Knowledge About A. mellifera
43
Honeybees expanded to North America with human-assisted migration during the
seventeenth century. Many Europeans fleeing wars, poverty, land laws or religious
persecution brought extensive beekeeping skills to the USA during the next two
centuries. Meanwhile, English colonists took bees to New Zealand, Australia and
Tasmania, completing human-assisted migration of A. mellifera around the globe.
Beekeeping became commercially viable during the nineteenth century with four
inventions: the movable-frame hive, the smoker, the comb foundation maker and
the honey extractor. These inventions still support commercial apiculture. A fifth
invention, a queen grafting tool, allows beekeepers to control genetic lines.
Honeybees are such efficient pollinators that industrialized countries developed
specialized agriculture dependent upon migratory pollination with honeybee, A. mellifera. The US Congress passed a Honey Bee Restriction Act in 1922, to protect A.
mellifera from the damage that tracheal mites were doing to honeybees in Europe,
until tracheal and varroa mites arrived in the 1980s, resulting in 50–80 % loss of their
colonies.
Different honeybee races can clash with pre-existing insect species. In the 1950s,
the honeybee A. mellifera scutellata (one type of African honeybee) was taken to
Brazil via human assistance, creating ramifications for the endemic bee species in
both South and North America. Similarly, A. mellifera was introduced to India and
China, but it competes with the smaller A. florea for floral sources.
Honeybees can adapt to minor changes in global warming, but colony collapse disorder is the most recent bittersweet reminder that human society threatens honeybee
habitats and breeding patterns on a global scale. Promoting genetic diversity of honeybees and providing safe environments are crucial steps towards future sustainable
agriculture.
2.3
Knowledge About A. mellifera
A certain amount of knowledge about A. mellifera in ancient Egypt is attested by
depictions of the bee and of hive beekeeping found during excavations. Statements
about honey and beeswax survive from Egypt, Greece and Rome, but most of them
record trade in these goods, or their use as offerings, gifts or the payment of dues.
2.3.1 Ancient Egypt
From about 3100 bc, the profile of the worker honeybee (A. mellifera) was used as
a hieroglyph in the topographical symbol of ancient Egypt. The earliest examples
excavated show four of the six legs and two of the four wings. The bee’s head,
thorax and banded abdomen were demarcated, as well as the two antennae and four
legs. Hives of bees were portrayed by around 2400 bc. Four early representations
of honey harvesting from hives have been found during the excavations in Egypt
44
2 Historical Perspective
Fig. 2.1 An early
representation of a honeybee:
a stone bas-relief from the
Sun Temple of Ne-user-re,
Abu Ghorab, Lower Egypt,
c. 2400 bc (drawing:
Egyptian National Museum
Catalogue). (Crane 2004)
(Crane and Graham 1985). One of them, reproduced in Fig. 2.1, shows that smoke
was used to pacify bees. Also, as combs taken from the hive were round in shape,
the beekeepers who produced them must have known how to get the bees to build
their combs directly across the horizontal cylindrical hives shown. The beekeepers
also probably understood—as traditional beekeepers do today—that the bees build
their combs in a nest cavity or hive at a constant separation. Thus, if two or more
guide combs are inserted in an empty hive, they must be separated at the bees’ comb
spacing. Traditional beekeepers in Greece use the combined width of the first two
fingers, or the length of the distal (outer) thumb segment, to determine the distance
between the midribs of adjacent combs. Although no written descriptions of bees or
beekeeping are known from ancient Egypt, the depictions suggest that beekeeping
methods reached a higher level there than anywhere else at the same period (2400–
1400 bc). The method used by traditional beekeepers in Upper Egypt today is similar
to that depicted in 1450 bc (Fig. 2.2). The method was also transmitted westward
along the north African coast and to Sicily, and some, but not all, parts of it reached
Greece and Rome.
2.3 Knowledge About A. mellifera
45
Fig. 2.2 Part of the
wall-painting in Rekhmire’s
tomb (no. 100), West Bank,
Luxor, Upper Egypt, c. 1450
bc (Davies 1944; Crane 2004)
2.3.2 Ancient Greece
In ancient Greece, it was known that each colony contained one larger bee, which
was regarded as the ruler and assumed to be male. Early Greek texts have praised
the large bee in the hive for its outstanding leadership abilities and wisdom. Some
of the texts praised the bee’s feminine gender characteristics and others praised its
masculine ones—using almost identical terms. In either case, all the other bees were
subject to their leader, from whom they did not want to be separated. These authors
concluded that praising bees was a common rhetorical device of classical Greek
writers, who were not concerned with the gender of the bees’ ruler. Writings on bees,
which survive from ancient Greece, contain a number of important statements. They
describe behavioural characteristics of A. mellifera more or less correctly, although
the physiological basis for these was usually not understood. Books I to VIII of
Historia animalium were written by Aristotle (384–322 bc), but the large amount of
information on bees is mostly in Book IX, whose author is unknown. The following
statements are quoted or summarized from Book IX, which state that outside the
colony, bees visit flowers, but only one kind on the same trip. They collect propolis,
‘the “tears” or exuding sap of trees . . . ’ and use it to ‘narrow . . . the entrances to the
hive if they are too wide’. ‘Others (of the foragers) carry water’ and bees ‘discharge
their excrement in flight’. When drones fly out ‘they soar up in the air in a stream,
whirling round and round’. Regarding swarming: ‘They say that, if a young swarm
goes astray, it will turn back upon its route, and by the aid of scent seek out its leader’.
‘In the colony, there is a division of work in the hive: some make wax, some make
honey, some make bee-bread (pollen), some shape and mould combs . . . .’. ‘Others
smooth and arrange combs’. Bees ‘store up another article of food resembling wax
in hardness, which by some is called sandarace, or bee bread. This they carry on
their legs’. ‘When the floral world is in full bloom then they make wax’ and when
bees are smoked they ‘devour the honey most ravenously’. They build cells ‘for the
Kings only when the brood of young is numerous’ and the ‘bees that die are removed
from the hive’. ‘When honey runs short they expel the drones’.
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2 Historical Perspective
In the 100s bc, Zenodorus of Sicily proved that, of the three regular figures, which
will completely fill an area, the hexagon (the cross-section of a bees’ cell) has the
greatest surface area (Betts 1921). From the Hellenistic Period (323–331 bc), the
concept of bugonia (birth from an ox) was prevalent; it was probably of Egyptian
origin. To produce a swarm of bees, an ox was to be beaten to death without breaking
the skin and the body to be enclosed with herbs in a special building for 9 days, after
which a swarm of bees would appear. The idea probably arose from a confusion
between the honeybee and the drone fly Eristalis tenax, and between larvae of the
honeybee and of the blow fly (Calliphora spp.). Other misconceptions have been
listed elsewhere (Crane 1999), many of them could have been corrected by quite
simple observations. Davies and Kathirithamby (1986) included a full discussion of
knowledge about insects in ancient Greece, and of the attitude to studies of such
small animals.
2.3.3 Ancient Rome
Roman writings include many statements similar to those in Greek texts (Crane 1990)
which supports the view that Greece was the source of Rome’s knowledge of bees.
Roman authors wrote much about bees and beekeeping, and the main passages by the
following authors such as: (1) Varro (116–27 bc) Res rusticae, Book III.16.1–382;
(2) Virgil (70–19 bc) Georgics, Book IV; (3) Columella (c. 60 ad) De re rustica,
Book IX.2–16; (4) Pliny (the Elder) (23–79 ad) Naturalis historia, Book XI, 4–16;
XXI, some of 43–49; (5) Aelian (died c. 220 ad) De natura animalium, scattered
references; and (6) Palladius (300s ad) Opus agriculturae, scattered references were
quoted by Crane (1994) with short commentaries.
2.4
Knowledge About A. mellifera up to 1500–1600
A number of the writings about bees from other Mediterranean civilizations have been
lost, but several were translated and preserved by Arab scholars who lived in Spain
during the period between the invasion by Muslims in 711 and their expulsion in 1492.
Some writers who lived between the 900s and 1100s were especially important in
preserving knowledge about bees, and in adding new knowledge (Monferrer 1991).
They include the following: Ibn Sina (980–1037) or Avicenna, who was born near
Bokhara in Uzbekistan and died in Persia; Ibn Wafid (born in 1008) of Toledo, who
studied in Córdoba; Abu Zacaria (1000s–1100) in Tunis; Ibn-al-Awam of Seville;
and Ibn Ruoshd (1126–1198) or Averroës, born in Córdoba.
Avicenna (unlike Aristotle) knew that ‘kings’ were reared in extra large cells.
Ibn-al-Awam stated that the smallest bees in the hive are females, which have a
sting. Larger bees are males, which take no part in the preparation of honey. The
‘kings’ are twice as large as the bees that make honey, and Ibn-al-Awam knew that
2.5 New Knowledge About A. mellifera between 1630 and 1800
47
it was advantageous to the beekeeper to have only a small number of these in a hive
(Bee World 1932). Many Greek writings were translated into Arabic; from these,
Latin translations were made and later disseminated widely in Europe. Books were
printed in Europe from 1459 onwards. In 1513, Gabriel Alonso de Herrera in Spain
published a compilation of writings on agriculture by previous authors (Alonso De
Herrera 1513) and Volume 5 of this work was devoted to bees. It reported what Greek
and Roman authors had written.
In 1586, a book by Luiz Méndez de Torres in Spain contained the following
explicit statement: . . . la aveja, que dizen maessa, o maestra, sin ayuntamiento de
macho, y sin dolor, echa de si una semilla, de que se engendran tres generos de avejas,
que son, maestras, y zanganos, y avejas. De suerte, que siendo la simiente una misma,
por razon de la diversidad de los vasos donde se pone . . . (. . . the bee, called the
maessa or maestro (mistress), without coupling with a male (this is incorrect) and
without the pain of childbirth, produces a seed from which are engendered three
kinds of bee—maestras, drones and ordinary bees—according to the different cells
in which the seed is placed. . . ) (Méndez 1586).
Meanwhile, in 1568 in Silesia, Nickel Jacob had published a book on beekeeping
(Jacob 1568) which included two significant new observations: (a) a colony with (or
given) young worker brood or eggs can rear a new Weisel (a masculine noun used for
the ruler of the colony and (b) when a hive of bees is put in a new place, the bees learn
its location by circling in the air above it. Important advances followed after Galileo
(1564–1642), in Italy, developed a compound microscope; he described it in 1610,
although the word microscope was not in use until 1624. But before that, Giovanni
Rucellai (1475–1525, also in Italy) wrote a poem Leapi—not published until 1539—
which described what he had seen of the bee’s external morphology, including the
proboscis and sting, by using a concave mirror (specchio lucido escavate). Rucellai
referred to the ruler as king (re), never as queen (regina). Galileo was a member of
a small but active scientific society in Rome, the Accademia dei Lincei (Academy
of the Lynxes) (Carutti 1883). In 1624, he gave a microscope to Prince Federigo
Cesi, the founder of the Academy, who used it to draw honeybees on a broadsheet
to be presented to the Pope (Fig. 2.3); they were the first insects to be depicted as
seen under a microscope (Crane 1963). Cesi also started to edit his large drawings to
produce a textbook on bees, Apiarium, which was to be indexed and made suitable for
quick reference (Alessandrini 1956). He cut up the text, stuck each item on a separate
sheet of a notebook and wrote notes and additions in the margins. His draft can still
be seen in the Academy in Rome, but Cesi died before he could complete the book.
2.5
2.5.1
New Knowledge About A. mellifera between 1630 and 1800
Observation Hives
In 1654 in England, John Evelyn was shown ‘Transparent Apiaries’ in the garden of
Dr. Wilkins at Wadham College, Oxford, although the ‘glass hive’ he drew appears
to have only small windows (Smith 1965). The diary of Samuel Pepys for 5 May
48
2 Historical Perspective
Fig. 2.3 Broadsheet
(100 cm × 64 cm) presented
to Pope Urban VIII in 1625,
with the first drawings of bees
made using a microscope.
(Crane 2004)
1665 refers to bees ‘being hived in glass (so that) you may see the Bees making their
honey and combs mightily pleasantly’. By about 1670, large sheets of glass were
available in England, and observation hives could then be made with glass sides. The
use of such hives for studying the activities of bees seems to have developed rather
slowly, although Réaumur (1740) believed that Swammerdam must have used one
in his studies of bees between 1669 and 1673. Robert Boyle, an Irish chemist and
physicist, said in 1688 that he had in his closet ‘a Transparent Hive, whence there
was a free passage (for the bees) into a Neighbouring Garden’ (Boyle 1688). In 1712,
the Italian astronomer Maraldi referred to his observations using a large number of
glass hives in a garden in Paris (Maraldi 1712), and in the same year, Joseph Warder
in England reported keeping bees in ‘transparent boxes’ (Warder 1712). In 1788,
Spitzner described his observation hive, which held a single comb between two large
sheets of glass, and he stated that returning foragers danced on the comb surface
(Spitzner 1788).
2.6
Reproduction and Gender in Honeybees
The statement by Luis Méndez de Torres in Spain in 1586, that the ruler bee was
female and the mother of all the other bees in her colony, slowly filtered through to
other countries. In 1637 in England, Richard Remnant wrote that ‘the females (from
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