The Biogeography of ..Apis mellifera..............................
 

by Kristen Coker, student in Geography 316

Thank you for visiting our site. This web pages was written by a student in Geography 316: Biogeography and edited by the instructor, Barbara Holzman, PhD.  All photos and maps are posted with specific copyright permission for the express use of education on these web pages. The students have tried to be as accurate as possible with the information provided and sources and references are cited at the end of each page.

Species Name: Apis mellifera

Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Apidae
Genus: Apis
Species: Apis mellifera
 

Description of Species:
   
  Photo of Apis mellifera taken by James H. Cane, USDA-ARS
 
 Biogeography of the Honeybee

            The common name of my organism is the Honeybee.  Its taxonomic

classification is as follows:

 There are hundreds of bee species around the world.  There are even more

subspecies that have been identified, while others wait to be identified. 

These honeybees can be found from the tropics to the Arctic Circle.  They

live in unique and highly specialized colonies.  Their evolution has divided

the species greatly, and largely affected the distribution of the Apis

mellifera.

            The Apis mellifera has the morphological characteristics common in the

insect class.  Its body is made with an exoskeleton of segmented body parts.

  They are the head, the thorax, and the abdomen.  They have 2 antennae, 2

compound eyes, 3 simple eyes, 6 legs,  and 2 pair of wings (on the thorax). 

They are generally 15-25mm in length however their specific sizes vary

greatly depending upon the species.  Their oval shaped bodies are distinctly

marked with yellow and black stripes of hairy skin. The bodies of the queen

bees are generally bulkier than the male bees.

            The Apis mellifera‚s habitat occurs in dry and sheltered places.  The queen

bee designates the living location for the hive.  The most important

elements in selecting a home is its proximity to nourishment, and its safety

from disruption.  Locations include inside hollowed tree trunks, tree

branches, previously inhabited bird nests, thatched roofs, under floorboards

of outhouses, or any protected nook.  Many queens prefer to live in

underground tunnels with varying preferences in how deep and long the

tunnels are.

            The distribution and range of Apis mellifera is quite large.  Various

subspecies can be found throughout the world.  Their location ranges from

Europe, to Asia, to North and South America, and even the Arctic Circle. 

Apis mellifera tend to live in temperate to warm climates, like the

Mediterranean and deserts.  They typically appear in the Northern climate

zones and can live from high mountain ranges to low and dense tropics.

            The mating routine of Apis mellifera is quick and calculated.  The entire

structure of the colony is based upon the needs of the queen to reproduce,

and to reproduce efficiently.  The drone bees (whose only purpose is to mate

with the queen) generally will fly around in a pattern above the ground

(50-100 meters).  Often times these flight patterns are re-used through the

years.  The queen bee gives off an odor, the drone bee responds and seeks

her out on his flying journey.  At about 1 meter he can see the queen. 

After inspecting her with satisfactory results, he will mate with her.  „...

copulation is violent and soon completed.  The drone‚s genitalia are

explosively exerted into the sting chamber of the queen; he is unable to

withdraw them and they are left behind when he literally „tears‰ himself

away.(Free 1982)  The male drone will die shortly after.  The queen can

continue to mate with other drones if her sperm storage sac (spermatheca)

has insufficient sperm.  This can continue for a few days with up to ten

drones in a relatively small amount of time.  The drone however will only

mate once, successfully.  There is significantly little available research

information on the mating habits of Apis mellifera for it occurs while they

hover above ground.  Researchers are unsure what attracts a drone to a queen

in time for mating, for within the nest a drone is not attracted to a queen,

whether virgin or previously mated(Kearns &Thomson 2001).  Once satisfied

with her sperm inventory, the queen will not need to copulate again for the

rest of her life.  She will have enough sperm stored to last her a lifetime

of egg laying.  The egg laying process generally occurs in the spring and

summer months so as to produce new workers in time for pollination and

nectar collecting.  The queen bee lays one egg in each cell of the hive(or

comb).  The eggs will be fertilized or unfertilized.  The unfertilized eggs

are drones while the fertilized eggs can be workers or queens.   Other

female bees will only survive and be able to reproduce when the queen bee is

dying or dead, and the workers are not subdued by the pheromones put out by

the queen.  In general, it takes about 24 days for a drone to develop from

an egg, 21 days for a worker, and 16 days for a queen bee.

            The different bees in a hive are all highly evolved and specialized.  The

queen Apis mellifera for example, has evolved so that she can only produce

eggs.  She is not capable of feeding herself, or her offspring, she cannot

produce wax, nor can she collect pollen or nectar.  The drones too are only

capable of mating with the queen.  They are useless when it comes to

foraging or building hives.  The worker bees compliment the system by doing

just what their name implies.  They conduct all of the work taking place to

ensure the success of the nest as well as the queen bee.  They build the

nest, produce wax, collect, deliver and store nourishment.  They also

procure the necessary ingredients to produce honey.

             Apis mellifera is significant to humans for many reasons.  Arguably its

most important trait is its practice of pollinating flowers and plants. 

Bees are keystone mutualists  in helping to propagate several species of

plants.  It can only be estimated how many species the bees have helped to

expand into different areas.  Along that same line, the bees have brought

with their plants all the organisms dependent on those plants, further

propagating plant and animal life.  Besides helping to pollinate and

increase plant life, the Apis mellifera has brought to humans their sweet

nectar, honey.  Humans have enjoyed eating honey and using its medicinal

qualities for centuries.  Also the wax produced by bees can be manipulated

for use in everyday products.  The products made by bees have been quite

beneficial to humans for years, and there are numerous successful beekeepers

around the world who take advantage of their gifts.

Evolution:

Evolution of Apis mellifera

            The evolution of Apis mellifera  is believed to have begun in the Cenozoic

era.  The first recorded fossils of Apis meliponoides  (later classified as

a separate genus, Electrapis) are an ancestral link to all recent Apidae.  

It is difficult for researchers to clearly delineate the phylogeny due to

irregular distribution of original apoid and derived characteristics among

subfamilies (Ruttner 1988).  It was found that there was an even

distribution of derived characters among the three groups of Apis  , though,

with the help of Winston and Michener developing a list of characteristics

of the Apis species.(1977)  These distinctions included both physical and

behavioral characteristics.  After seeing the pattern among the Apis , a

cladogram was designed to incorporate the Apinae to Bombinae, moving from

Meliponinae(Winston and Michener 1977).  It is apparent that all subfamilies

of Apidae are a  descent from a common gene pool in the Eocene, while

Meliponinae is a little older.  (Ruttner 1988).

  A phylogenetic tree based on information acquired from the Tree of Life Web Project.

            It is believed that each subfamily radiated at separate times.  The
Meliponinae in the early Tertiary(Wille 1980), the Bombinae in late Tertiary
(Williams 1985), and the Apinae in the end of the Tertiary-Pleistocene.(Ruttner 1988)  The Meliponinae have 18 genera, 500 species; Bombini has 3 genera, about 290 species; Apini 1 genus 4 or 5 species, but with many subspecies.  Due in part to the time these species have had to evolve, and their consequent number of taxa the subfamilies of Apidae have developed uniquely to their environments (Ruttner 1988).  While the Meliponinae has larger perennial colonies, the Bombini have smaller seasonal colonies (often in cold, even arctic climates), and the Apini has adapted to many different environments allowing for different successful
locations.

            It is hypothesized that the two species A. cerana  and A. mellifera  are

still in the beginning states of speciation (Ruttner and Maul 1983).  This

hypotheses would indicate a splitting before or during the Pleistocene which

in turn indicates that they were isolated during the last glaciation.  Their

distribution into the temperate zone proves a postglacial pattern-therefore

they have existed for only about 50,000 years. (Ruttner 1988)

            Arguably the most important change or adaptation in the physiology and

behavior in the Apinae family is the development of multi-comb nesting and

thermoregulation.  These developments allowed the Apis to adapt to diverse

climates and led to the major diversification of species and sub-species. 

Researchers have had a difficult time tracing the steps back to the Miocene

for the course of the Apis  evolution has been punctured with long term

stagnation and then rapid evolution.  Ruttner states, „. . . . the

phylogenetic tree of Apinae is a bare structure with a few incipient

branches at the extremity . . . . „(pp.30)

            It is hypothesized that in the early Miocene species of Apis could have

survived in central Europe, but not into the Pliocene.  When the climate

became too cold, the Apis disappeared from Europe and survived in tropical

South Asia.  It is believed that sometime in the Pliocene or early

Pleistocene the Apis began to evolve into a state which allowed for thermal

homeostasis.(Ruttner 1988)  This allowed for the Apis species to be

independent of their environment.  It allowed them to live in extremely

diverse climates.  This in turn gave way to new behaviors including

multicomb nesting, and the ability to survive for months of cold weather in

a hibernation mode.  The success of these adaptations led to the overall

increase in Apis territory and species diversification.

Distribution

Distribution of Apis mellifera

            There exist four species under the title Honeybee, or Apis.  The most

common and well researched is Apis mellifera.  It is native to Africa,

Europe and Asia, however it is widely distributed around the world.  Early

pioneers brought it to the Americas and to Australasia because of its high

honey yield. (Free.1982)  The other three species-A.cerana, A. dorsata, A.

florea  are only found in southeastern Asia.  They are differentiated by few

physical qualitative characteristics as well as behavioral characteristics. 

The family Apidae, and specifically the genus Apis  are a source of constant

aggravation for taxonomists.  There are several thousand species and within

a single group there can be dramatic differences in the size of the anatomy,

making it difficult to determine whether the particular specimen is of one

species or another.  This is why behavioral characteristics count as a

significant factor in determining taxonomy.

            It is believed that the fossils which have been recovered from Baltic amber

of many Hymenoptera are the link to the first Apis mellifera.  They have

been recorded as being from the upper Eocene in the early Cenozoic period. 

Although no Apis  were found, a record of a species of recent meliponine

genus, Trigona  was.(Kelner-Pilault 1969; Zeuner et al. 1976)  Apis

meliponoides  classified by Buttel-Reepen(1906) was later classified as a

separate genus Electrapis, (Cockerall 1908) but is of significant value for

it demonstrates several variable, intermediate characteristics of all three

subfamilies of Apidae (including Apis and Bombus).  It appears that the

Electrapis  was a group of species in an ancestral line to all recent

Apidae.(Ruttner, 1988)

            Quality specimens of a true Apis  have come from the lower Miocene and were

found in Rott, Germany.  From the upper Miocene are fossils of the Randecker

Maar and Bottingen (Germany).  They are both classified as Apis armbrusteri 

with numerous subspecies(Zeuner et al.1976).  Found in East African copal,

fossilized honeybees were found that do not differ from recent A. mellifera.

  They were thought to be of the Pleistocene era, but are of very recent

origin. (Burleigh and Whalley1983).  It is believed that an Apis species

could have lived in Europe during the late Miocene and then during the

Pliocene migrated toward the Meditteranean basin due to climate

changes(Ruttner 1988).  As incredible evolution took place over several

thousands of years, the Apis  became adapted to several climates and began

to spread to Africa and southern Asia.

            Throughout the world one could find Apis mellifera  as well as numerous

other species of bees.  Bees are found in all climates, at all altitudes,

and in several different environments be it thick bushes, underground, above

ground in trees, or in man-made hives.  A. mellifera  can be found in the

tropics of Africa and S. America, India, throughout Europe, China, N.

America and Canada, as well as the Arctic circle.  Globally there are

approximately 25,000 named species of bees, with an estimated 40,000 species

existing in total.  In the United States alone there are about 5,000

species. (Buchanan, 2002).  The enormous distribution of A. mellifera  is

due mainly to mankind.  Man has been breeding bees for all of history in

quest of honey.  This desire for honey has been the largest contributor than

any other force of the widespread distribution of A. mellifera.  This also

has accelerated the quantity of morphological differences among bees,

ultimately producing the thousands of species and subspecies of A.

mellifera.

Map of Distribution:
 
   

Other interesting issues:
   

Bibliography
(for web sites put in the url)
 

send comments to bholzman@sfsu.edu
 

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