Solitary Bees

Author(s): Suzanne W. T. Batra

Source: Scientific American, Vol. 250, No. 2 (February 1984), pp. 120-127
Published by: Scientific American, a division of Nature America, Inc.
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 Solitary Bees
Most species of bees are not social like the honeybee. They are

solitary, meaning that the female buJids her own nest. They play

a profoundly important role in pollinating crops and wild plants

by Suzanne W. T. Batra

or most people the beehive and Oxaeidae and Fideliidae, which are an environment shown to be satisfacto­

F the intricate social organization of

the honeybees that populate it are
small families with a limited distribu­
tion and no common names, and the
ry by the survival of her ancestors.
Bees that nest in the ground usually
the hallmark of bee life. In reality more Apidae, or honeybees, stingless bees have plenty of room for nest sites. They
than 85 percent of the some 20,000 bee and orchid bees, all of which are most­ face, however, the problem of protect­
species are not social but solitary. Each ly tropical, and bumblebees, which are ing for almost a year the nutritious hon­
female independently mates, makes her mainly in the Northern Hemisphere and ey-and-pollen provision they have laid
own nest of about 10 brood cells, stocks are the most able of all the bees to toler­ up for their brood (and the delicate,
the cells with food for the young, lays an ate low temperatures. helpless, grublike brood itself) from soil
egg in each cell and dies before the next moisture and a wide variety of soil or­
generation emerges. The solitary bees he nests and brood cells of most sol­ ganisms such as bacteria, yeasts, fungi,
play immensely important roles in eco­ T itary bees are made underground. nematodes and mites. The large majori­
logical systems, particularly in the polli­ Bare, dry, light soil exposed to the morn­ ty of ground-dwelling bees line their un­
nation of crops and wild plants. Indeed, ing sun is preferred. In such habitats derground brood cells with special wa­
farmers in some parts of the world are many nests of their relatives, the ants terproof secretions before provisioning
beginning to make specific arrange­ and the solitary wasps, may also be them. These bees possess an abdominal
ments to accommodate solitary bees for found. In a favorable area thousands of gland (Dufour's gland) that opens near
the pollination of crops (such as alfalfa) nests of ground-dwelling solitary bees the sting. In some species it occupies
that are not pollinated effectively by may form dense aggregations. The fe­ half of the abdominal cavity. The gland
honeybees. males do not, however, cooperate In contains an oily liquid of a fragrant-to­
Bees apparently evolved from pre­ nest-building chores. musky odor. (A few species collect plant
daceous ancestors such as the solitary The largest recorded aggregation of resin or cut pieces of leaf to line their
mud-dauber wasps in about the middle nests occupied about 360,000 square brood cells. The Fideliidae and other
of the Cretaceous period (100 million meters extending for seven kilometers bees that live only in deserts do not line
years ago), when flowering plants be­ along a bank of the Barysh River in the their cells at all.)
came the dominant vegetation on the U.S.S.R. It consisted of an estimated 12 By inducing bees to nest in narrow,
earth. Now bees live everywhere, show­ million nests made by two species of glass-sided chambers filled with soil I
ing the greatest abundance and diversity sweat bee and a species of miner bee. In was able to observe their subterranean
of species in semiarid, warm temperate about mid-May every year I receive at nest-making behavior. After a chamber
climates. Many solitary bees do not my U.S. Department of Agriculture of­ for a cell is excavated by the bee and
resemble honeybees in general appear­ fice in Beltsville, Md., many agitated tel­ often polished to a shiny smoothness by
ance but instead look more like wasps, ephone calls about swarms of solitary the pygidial plate at the end of the abdo­
flies or bumblebees. They range in bees nesting on lawns in the Washington men, dollops of Dufour's-gland secre­
length from 1.5 to 46 millimeters. Some area. These are usually the membrane tion are deposited in it and are repeated­
species are hairless and shiny, others are bee Colletes thoracicus, which is active as ly smeared around by rhythmic sweeps
densely furry. Although many of them an adult for about six weeks. The largest of the bee's tongue, by special velvet­
are drab, some (such as the emerald aggregation of this honeybee-size insect like pads on the abdomen or by hairy
green orchid bees) are quite beautiful. included about 104,000 nests in a subur­ brushes on the legs. Most solitary bees
The hair or the spots and stripes may ban backyard of 1,187 square meters. fly about during the day collecting pro­
include all the colors of the rainbow. Aggregations of nests may persist for visions, lay an egg in the afternoon, seal
Solitary bees are distributed among many years. For example, a species of the cell, begin digging a new cell cham­
nine families: the Colletidae, or mem­ miner bee has occupied an adobe wall ber in the evening and apply the Du­
brane bees, which are most numer­ near Baltimore for at least 40 years. The four's-gland secretion at night, so that
ous and diverse in the Southern Hemi­ aggregations are probably maintained the new cell is dry and ready for provi­
sphere; the Andrenidae, or digger bees, because the young females on their first sioning the next morning. Such bees
which are mainly in the Northern Hemi­ flight memorize visual landmarks in the work day and night with little respite.
sphere; the Halictidae, or sweat bees (so area, just as new honeybee workers re­ Henry M. Fales and Abraham Hefetz
named because they are attracted by hu­ member the location of their hive. Pher­ of the National Heart, Lung, and Blood
man sweat), of worldwide distribution; omones (chemical signals) or odors re­ Institute and I analyzed the secretions of
the Megachilidae, or leafcutter and ma­ leased by other females in the aggre­ Colletes membrane bees. We found that
son bees, also worldwide; the Antho­ gation may also be attractive. After the Dufour's gland contains fragrant,
phoridae, or carpenter and miner bees, mating, a young bee returns to make musky-smelling compounds (macro­
predominantly tropical; the Melittidae, her own nest among the other nests in cyclic lactones). When they are deposit-

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ed in the brood cell, they polymerize to For comparison we also analyzed the rasp away and eat the cell lining. Until
form a flexible, transparent, waterproof Dufour's-gland secretion and brood-cell the miner-bee larvae's feeding habits
membrane composed of a natural poly­ lining of Anthophora abrupta, a miner were discovered the honeybee was the
ester (mainly 18-hydroxyoctadecanoic bee. This bee's large Dufour's gland only bee known to feed its larvae with
acid and 20-hydroxyeicosanoic acid). holds a mixture of transparent, slightly a secretion (royal jelly, secreted orally
The membrane forms a miniature plas­ fragrant oily triglycerides. When they and consisting mainly of protein).
tic bag in each cell. The bag even has a are secreted into the brood cell, they

B
"zip lock" closure: a flap that is lifted are converted, possibly by salivary en­ ees that do not live underground
and sealed by the female bee after she zymes, into white, waxy diglycerides need not protect their young so
has laid an egg. The membrane is quite with a rancid odor. This secretion is also carefully from excess moisture, and
durab1e, remaining intact in soil for a added to the provision for the larvae. probably for this reason they have rela­
year or more. The developing larvae eat it and even tively small Dufour's glands. Most ma-

k�
SOLITARY BEES are compared with the honeybee, Apis mellifera gil/iea ( f ); the miner bee, AI/lhophora abrupla (g); the orchid bee, EIl­
(a), in drawings that are done to a common scale. The solitary glossa igl/ila (h),shown with its unusually long tongue extended; a mason
bees are the alfalfa leafcutter, Megachile rollil/dala (b); the membrane bee of the genus AI/lhidillm (i); a digger bee, AI/drel/a carlil/i (j), and
bee, Colle/es Ihoraciclls (c); the alkali bee, Nomia JIIelal/deri (d); a a silk-making membrane bee of the genus Hylaells (k). In life Hylaells
cuckoo bee, Nomada Illteoloides (e), so named because it lays its eggs is four millimeters long. The bees portrayed represent only a few
in the nests of other solitary bees; the carpenter bee, Xylocopa rir- of the many solitary-bee species, of which there are some 17,000.

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son bees and leafcutter bees use existing The secretion of silk by adult insects is These microorganisms, usually present
aboveground holes as nests, building quite rare. in the nectar from flowers, are normal­
brood cells of mud, resin and plant fiber Most female solitary bees, like the ly deposited along with the provision.
or of neat layers of ovals and circles honeybees and the bumblebees, can de­ Their growth is prevented, however, .by
precisely cut out of leaves and petals. fend themselves individually by sting­ the high sugar concentration; with soli­
Leafcutters may damage plants. In an ing, but their venom is usually mild. The tary bees, as with honeybees, the excess
extreme example the yield of two hec­ nests of solitary bees, unless they are water in the honey evaporates away
tares of soybeans was reduced 30 per­ densely aggregated, seldom attract ver­ as the flower nectar is converted into
cent by the removal of leaf tissue by tebrate predators to their meager pro­ honey by enzymatic reactions. If yeasts
bees nesting nearby. Some mason bees visions and brood. Therefore the bees grow unchecked, the bee larva gets indi­
build clustered brood cells of mud or have not developed the aggressive mass­ gestion and becomes weak or dies. The
resin on branches or walls. One com­ stinging counterattack provoked when fermented provision is then overrun by
mon species of solitary bee is notable honeybee and bumblebee nests are dis­ filamentous fungi.
for defacing Egyptian and Greek monu­ turbed. Solitary bees' nests, however, Working at Utah State University
ments with its numerous large, lumpy are particularly vulnerable to attack by with two of my colleagues in the De­
nests of dried mud. other invertebrates and by microorgan­ partment of Agriculture (George E. Bo­
Carpenter bees drill neat round holes isms, and the bees have evolved ways to hart, an entomologist, and my husband
with their powerful mandibles into repel them. Lekh R. Batra, a mycologist), I found
wood, bamboo or pithy stems. In their Fungi are the primary enemies of soli­ that bee adults and broods are attacked
galleries they construct series of brood tary bees that nest underground. The by at least 124 species of fungi. The soli­
cells separated by partitions of aggluti­ bee must work quickly to finish the tary alkali bee has developed an inter·
nated sawdust. They sometimes do sig­ preparation of the provision and to lay esting method of protecting its brood
nificant structural damage to the beams an egg so that the larva can consume the from some of the 49 species of fungi that
or eaves of buildings. food before it spoils. The brood-cell lin­ afflict it. In common with most other
Membrane bees of the genus Hylaeus ing isolates the contents from free water solitary bees the alkali bee seals each
are wasplike in appearance and carry in the soil. Water vapor enters many cell after provisioning it and laying an
provisions internally as wasps do in­ cells, however, along with the gases egg. It has long been assumed that such
stead of on special hairs in the manner needed for the respiration of the larvae. "mass provisioning" bees have no fur­
of other bees. Hylaeus bees are unique in The provisions of most bees consist of ther contact with their young. When the
making brood cells of true silk inside a mixture of honey and pollen. The sug­ contents of a cell become infested with a
existing holes. The silk is secreted orally ars of the honey are hygroscopic (they fungus, however, the mother bee opens
and applied by the female bee's broad attract moisture from the air), and so the cell and packs soil into it. This proce­
tongue. Small predatory wasps belong­ they may gradually become more di­ dure reduces the amount of air available
ing to the family Pemphredoninae also lute. This may account for the fact that for the growth of the fungus and so pre­
(uniquely among wasps) make cells out bee larvae may weigh more than their vents the fungus from spreading to adja­
of silk. It is possible that bees have provision originally did. All is well with cent cells. Although some soil-dwelling
evolved, through primitive forms such this system unless populations of yeasts bees may annually lose as much as half
as Hylaeus, from this group of wasps. and bacteria in the provision explode. of their brood to fungi, the species that
nest in drier sites such as sound wood,
stems and adobe walls are usually less
severely affected.

S cuckoo bees or inquilines, are com­

pecialized solitary bees, known as

mon parasites of other solitary bees.

Like its bird namesake, a cuckoo bee
does not make its own nest but instead
lays its eggs in the nests of other bees.
About 15 percent of all bee species are
cuckoo bees. These parasites may be
quite host-specific, each attacking a sin­
gle host or a group of related host spe­
cies. Most families of bees include some
cuckoo species, which may attack hosts
belonging to their own family or to oth­
er families. The young larvae of many
cuckoo bees have long mandibles that
serve to crush the larva of the host bee;
older cuckoo-bee larvae, feeding after
the host has been eliminated, have nor­
mal small mandibles. Usually the most
easily observed cuckoo bees are the
slender, wasplike, rust-colored or yel­
OPENING OF low-and-black species of Nomada (An­
MANDIBULAR thophoridae), which flit about in early
GLAND
spring near the ground among the wood­
land flowers in search of temporarily
FEATURES OF THE ANATOMY of typical solitary bees include the pygidial plate, which
unoccupied nests of their Alldrella hosts.
the female uses to smooth the inside of a nest she has excavated in the ground, and Dufour's
gland (within the abdomen), with which she applies a secretion that waterproofs the inside of
A solitary bee must spend several
the nest. Mandibular-gland secretions serve in communication and self-defense. The scopa, hours daily away from its nest foraging
which in some species is on the abdomen, is an array of hairs for carrying pollen. A solitary for nectar and pollen. During this time
bee walks on its knees (the basitibial plates) when it is ex(avating or provisioning its burrow. the helpless brood is particularly vulner-

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able. The brood cells of some species are
tightly sealed with a thick cap of resin,
secretion or clay, but the cells of many
solitary bees have a permeable cap of
lightly packed soil, leaves or wood dust
that is readily penetrated by insect para­
sites and predators. The mother bee may
try to protect the nest in her absence by
positioning it so that the entrance is hid­
den, by temporarily filling the burrow
with soil, by heaping soil in a small tu­
mulus over the entrance or by building a
compact turret of earth that may deter
crawling insects. Nevertheless, solitary­
bee broods suffer loss<:s to a variety of
predatory and parasitic wasps, ants, sev­
eral kinds of flies and beetles, strepsip­
terons, mites and nematodes. Some of
the parasites deposit their eggs or larvae
on flowers, where they are accidentally
picked up by the foraging bee, brought
home to the nest and included in the
brood-cell provision, where they lie
ready to attack the bee larva.
The colorful, symmetrical and fra­
grant blossoms of higher plants evolved
to attract pollen-dispersing animal life.
Bees are the most efficient and effective
of all pollinators for several reasons, in­
cluding their sheer abundance, their rap­
id flight, their tendency to visit several
flowers of the same species in succes­
sion, their need for large quantities of POLLEN-CARRYING APPARATUS is the scopa, the group of branched hairs near the top

nectar and pollen and their specialized of this electron micrograph of the hind leg of a sweat bee of the genus Lasiog[ossul1I. Sever­
al spherical grains of pollen are visible in the scopa. The enlargement is about 62 diameters.
hairs, which can trap and hold as many
as 15,000 pollen grains per bee.
An individual bee may possess several
types of hair, which are grouped on the
body and legs according to their func­
tion. The dense hairs on the hind legs of
most solitary bees form a pollen-hold­
ing brush called the scopa. (In leafcut­
ter and mason bees it is below the
abdomen.) Honeybees and their rela­
tives have shiny bare patches on their
hind legs known as pollen baskets or
corbiculae, which carry nectar-moist­
ened lumps of pollen or nesting materi­
al. The hairs of bees are modified ac­
cording to the type of pollen collected
by the particular species. For holding
fine, dry pollen grains a cluster of down­
like, branched, plumed hairs is best.
Large or sticky pollen grains call for
stiffer hairs. Such hairs often have short­
er branches or are formed with spiral
ridges resembling a corkscrew. They
may also have spatulate or hooked tips.
Combs of bristly hairs on the legs and
mouthparts of bees serve for grooming
and for transferrring pollen to the scopa.
The Dufour's-gland secretion is applied
with special straight hairs in tufts resem­
bling a paintbrush. To collect floral per­
fumes or plant oils some solitary bees
have velvety hairs that form sponges
and flattened hairs arranged in combs.

T of wild plants and crops is incalcula­

he value of bees for the pollination
INNER SURFACE of the sweat bee's hind leg shows (left) the spiral-ridged hairs that serve
ble. No price can be placed on wild to manipulate pollen and (lower right) the penicillus, used somewhat like a paintbrush for
plants, which are pollinated primarily spreading a waterproof secretion in the nest. The enlargement of the hairs is 300 diameters.

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by solitary bees. It has been estimated gle crops are planted as monocultures, well-sprayed commercial orchard with
that in the U.S. the value of some 90 where all plants flower simultaneously the bee populations on flowering trees in
crops that depend on insect pollination for a short time, solitary-bee popula­ abandoned orchards or at the edge of a
for maximum yield is $17 billion per tions may be insufficient to handle .the forest. In order to achieve adeq uate pol­
year. If seeds of insect-pollinated forage pollination, and when the flowers finish lination in the absence of enough native
plants for livestock are included, about blooming, nothing is left for the bees to bees, growers have to rent hives of hon­
a third of the nation's food supply de­ eat. Cultivation and construction elim­ eybees from beekeepers at a cost of
pends directly or indirectly on pollina­ inate supplementary floral hosts and about $20 per hive. Hundreds or even
tion by insects, mainly bees. It is rea­ nesting areas. Irrigation may destroy thousands of hives may be needed. The
sonable to assume that solitary bees nests. It also increases the damage done value of pollination services far exceeds
account for at least half of this crop­ to the bee brood caused by fungi. The the value of the honey and wax obtained
pollination activity in the U.S. and per­ careless application of pesticides kills from honeybees.
haps more in areas where beekeep ing is many bees. The sulfur dioxide in pol­ Honeybees pollinate many types of
less prevalent. luted air can cause bees to reduce their plant, but some plants are pollinated
Unfortunately populations of soli­ flying time. only ineffectively. For example, the
tary bees decline under the conditions These effects can be observed by com­ flower of alfalfa has a little spring in it
of large-scale modern agriculture and paring, say, the bee populations on flow­ that is triggered when a bee pushes her
urbanization. When large fields of sin- ering apples in the center of a large, head into the flower. The stamens pop

NESTING ARRANGEMENTS of six species of solitary bee are de­ the period of several weeks when the adult female is active. The nest
picted. Each nest has several individual cells, each of which the fe­ of the alkali bee, Nomia melal/deri (a), is made in moist soil. The mem­
male supplies with provisions for the larva before she lays an egg. brane bee, Co{{eles·lhoraciclIs (b), which also nests in soil, gets its com­
The cells are usually made at a rate of one a day (in good weather) in mon name from the fact that it lines the inside of each cell with a poly-

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up and rap the bee on the head, dust­ mind being rapped by the flowers, and cially prepared soil that is maintained at
ing, her with pollen. Honeybees appar­ they collect the pollen for provisioning just the right level of moisture to attract
ently do not like being rapped on the their nests as it is showered on them. The and hold dense aggregations of alkali
head, and since they are fairly intelli­ alfalfa-leafcutter bee, originally Euro­ bees. There may be as many as 2,100
gent insects, they learn to poke their pean, quickly visits many flowers in suc­ nests per square meter.
long tongue into the side of the flower, cession and is a highly efficient pollina­ Another solitary bee utilized com­
avoiding the spring.,Thus they take nec­ tor. It nests in groups. Farmers make mercially is a mason bee, the Japanese
tar but accomplish no pollination. Be­ suitable holes in "bee boards," which hornfaced bee. Apple flowers are often
cause alfalfa is one of the major forage they put in shedlike shelters in the alfal­ not pollinated efficiently by honeybees,
legumes, the supply of seed (a $100- fa fields. The brood cells of this bee are which tend to work slowly, to avoid
million-per-year industry in the U.S.) built out of bits of alfalfa leaf. touching the stamens and to fly away to
is of critical importance. The alkali bee is a species of sweat bee forage on dandelions and other weeds.
native to the U.S. It likes to nest on alka­ Some 40 years ago Japanese farmers no­

I
n the northwestern U.S. and Canada line flats, where the subsoil stays fairly ticed that the little hornfaced bees nest­
two species of solitary bee-the alfal­ moist. Growers who accommodate this ing in their thatched roofs were ener­
fa-Ieafcutter bee and the alkali bee-are bee construct "bee beds," which consist getic and effective pollinators of apples.
intensively managed to pollinate alfalfa. of plastic-lined trenches in the alfalfa Over the years the management of this
These persistent insects do not seem to fields. The trenches are filled with spe- bee has been perfected, and bee shelters
containing hollow reeds for nesting are
seen throughout the apple-growing dis­
tricts of Japan. Recently, in collabo­
ration with Yasuo Maeta of Shimane
University and other Japanese workers,
I have successfully established popula­
tions of this Asian pollinator in the east­
ern U.S., where it is being tested for
its adaptability to the climate and its
acceptance of the available nesting ma­
terial.
Many species of bees have life cycles
and habits that depend on the availabil­
ity of specific host plants. Bees that gath­
er pollen from one species or a few
closely related plants are termed oligo­
lectic. Bees such as the honeybee, which
can gather pollen from a variety of
SOIL PLUG plants because of the hive's long collec­
tive period of adult activity, are termed
polylectic.
Flowering plants and oligolectic bees
have developed some remarkable mu­
tualistic relations. For example, the
American native gourds are most effec­
tively pollinated by squash bees, which
are several native species of miner bee.
Some species of sweat bee pollinate eve­
ning primroses and fly at night, which is
unusual for bees. This activity is aided
by their eyes, which are enlarged for
light gathering. Other specialized miner
bees pollinate wild morning glories. Per­
haps the most highly specialized solitary
bees are those visiting flowers that yield
oils, which are collected instead of nec­
tar (for mixing with pollen in the provi­
sion for the brood); to pick up the oil
these bees have hairs arrayed in comb­
like structures.
Orchids are among the most special­
ized of flowers. Some of them have
evolved unique ways of exploiting the
behavior patterns of solitary bees. Some
orchids have false nectaries: colored
spots yielding no nectar. When a bee
probes a false nectary for nectar, a pol­
linium (pollen packet) is attached to the
insect, which carries it along to pollinate
the next orchid. This deception works
ester membrane of its own making. The miner bee, Allthophora abrupta (c), employs a differ­ until the bees learn that the nectaries
ent technique in dry soil. The mason bee of the genus
Chalicodoma (d) builds its nest on the yield nothing.
surface of wood or rock; the carpenter bee, Xylocopa "irgillica (e), drills into wood. Some Other orchids have color patterns and
growers of alfalfa make grooved "bee boards" (J) for alfalfa-leafcutter nests to aid pollination. aphrodisiac odors that resemble the ap-

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pearance or the odor of female bees or Orchid bees apparently respond to species of carpenter bee and mason bee
wasps. Males are attracted. They try to other odors. Males of one species have the males are aggressive and battle to
copulate with the flower and in so doing become a nuisance because they are at­ exclude other male bees from their
pick up a pollinium, which is deposited tracted to houses that have been dusted small home territory.
at the next enticing orchid. Still other with DDT for mosquito control. They In ancient India the territorial bat­
orchids mimic certain male miner bees. collect the powder and pack it into their tles of carpenter bees were called "bee
When an aggressively territorial male leg slits. The compound does not seem sport." It is said that the homing of such
bee sees the orchid flower nodding in the to harm the male bees, but whether or bees was exploited for the carrying of
wind, the bee strikes it in an effort to not it has the intended aphrodisiac effect tiny messages. Much of the carpenter­
drive the "rival" away. As a result a pol­ on the females is not known. bee activity noticed by homeowners to­
linium is stuck to the bee's head. day consists of males zooming after one

M
The most complex relation is the one ale solitary bees ordinarily emerge another and even heading in the direc­
between certain South American or­ from the pupa a few days earlier tion of passing birds and airplanes. The
chids that yield neither nectar nor pollen than the females of their species. They territorial males stage battles of bluff,
and several species of orchid bee. The feed on floral nectar and pollen and zigging and zagging face to face until
male bees have swollen hind legs with search for newly emerged virgin fe­ one of them gives up and leaves.
hair-lined slits leading to internal pock­ males. In some species the males form Some Anthidium mason bees are un­
ets and have velvety pads on their front loose groups that jointly patrol the nest­ usual among insects because the males
feet. The males form leks, or all-male ing site, nearby landmarks or flowers are larger than the females. The reason
swarms, that attract females for mating. that are regularly visited by females. is probably that the males butt and grap­
Certain males are more attractive than When males see a female bee or an ob­ ple with each other in their aerial bat­
others because they have collected mix­ ject shaped like one, they pounce on it, tles; physical strength and weight would
tures of fragrances from the orchids. but unless a sex pheromone is present make it easier to retain a territory visited
They brush the flower's odor glands they let it go immediately. If a virgin by females. These males attack other in­
with their front feet and stuff fragrant female appears, a pile of writhing males, sects that wander into their territory.
particles into their hind-leg slits. While all attempting to mate, may form rapid­ They can break the intruder's wings with
the bee is doing so it picks up a pollin­ ly on top of her. Usually the first male to their strong mandibles and puncture its
ium from the orchid. Since these bees grasp the female holds on to her tight­ body with their stout abdominal spines.
can fly as far as 23 kilometers in a day, ly and succeeds in mating. Males of Curiously, male bees that fight over ter­
they are effective pollinators for sparse­ most species merely compete to reach ritory during the day may cluster togeth­
ly distributed plants, such as certain of the females first and are not noticeably er peaceably in the evening as they sleep.
the orchids. aggressive toward one another. In some In 1802 the naturalist William Kirby

2 3 4 5 6

7 8 9 10 11

NEST-MAKING ACTIVITY of an alkali bee, Nomio me!ollderi, (4). Now she is ready to lay in provisions (5, 6), consisting of a honey-pol­
is followed over a period of 24 hours. At the outset (1) the female len mixture, which will sustain the larva. After grooming the ball of
has finished one cell and has begun excavating another. When she provisions (7) and removing debris from the cell (8) she does more
reaches the appropriate depth, she packs soil along the edges(2). Then polishing of the cell walls(9) and lays an egg (10). She caps the cell
she smooths the inside of the cell with her pygidial plate (3) and ap­ (1 J) and adds a plug of dirt that comes from the early stages of her
plies a waterproof lining with a secretion from her Dufour's gland (12).
excavation of what will be the third cell in the nest-building cycle

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wrote: "I have often thought that if gen­
tlemen, who amuse themselves with
chemical experiments, should direct
their attention to insects, it might lead

How to put the

to the discovery of some powerful med­
icines. The variety of strong scents,
which these little creatures emit, is won­

big board on a
derful. . . . Many Melittae have a strong
scent, in some approaching to that of
gariick or onion. " An interesting if

small screen.
somewhat risky pastime is to carefully
sniff captive live bees and wasps, which
indeed produce a spectrum of odors that
evidently serve in communication and
defense. Small glands at the base of the
mandibles release when the mandibles
are opened various combinations of ter­ To take stock of the situation, an investor
penes, aldehydes, ketones, esters, pyra­ could use the IBM Personal Computer.
zines and other pungent or fragrant With our Dow JonesT" Reporter, * a device
compounds. called a modemt and a telephone, you can access
Some male bees apply such secretions
Wall Street and the world.
to vegetation to delineate their territory.
In other species the secretions serve as Tap the Dow Jones
attractants for one or both sexes. For News (Retrieval Servicet for
example, if a large number of bees such historical and current stock
as species of Colletes, Nomia or Antho­ quotes-day or night.
phora are netted at an aggregation of
(Use our Dow Jones
nests, they bite the net in an effort to
escape, releasing mandibular-gland se­ Reporter not only for easy
cretions as they do so. Other bees down­ access, but to save money
wind are attracted toward the netful of on connect time.)
captive bees. That it is the mandibular­ Pull 10K extracts of
gland secretions and not something else
over 6,000 companies.
that is attracting the other bees has been
demonstrated by the fact that the bees Get industry news.
are also attracted by excised mandibular Even enjoy sports news
glands or by the appropriate synthetic when you've had your
compounds mixed in the right ratio. The fill of business.
small carpenter bees Ceratina and Pithi­
To better manage your
tis produce pungent mandibular-gland
secretions when they are annoyed. They portfolio, visit an authorized
smear them over their bodies and the IBM Personal Computer dealer. And
intruder. These secretions are repellent learn how a small IBM investment can give
to nest-invading ants. you a high yield in quality, power and
Social behavior among the bees prob­
ably arose because the survival of the
performance. -:--:-_ -=. ='"
- ---
- - ---

brood is enhanced if a guard bee is pres­ - - - ---

---- - -
--_.-

ent while the forager is away. Another

benefit is that two or more bees can
share the work of nest construction and
provisioning.
Such behavior could have arisen in
two ways. The first is that the mother
bee came to have a relatively long life
span, so that she could help her daugh­
ters and be helped by them; this pattern
is seen in some sweat bees and carpenter
bees. The second way is that bees of the
same generation came to share nests and
cooperate in nest making and provision­
ing; this pattern is seen in a few species
of sweat bees, digger bees, mason bees
and orchid bees. Truly social behavior,
with a division of labor in which some
bees are queens, dominating the others The IBM Personal Computer
A tool for modern times
and laying the eggs, and other bees are
nonreproductive workers, is relatively
rare among bees, being known only in
some sweat bees and some small carpen­
ter bees in addition to the more famil­
iar social stingless bees, bumblebees and For more information on where to buy the IBM Personal Computer. call 800·447·4700. In Alaska or Hawaii. 800-447-0890.
honeybees. tNot supplied by IBM. 'Dow Jones is a trademark of Dow Jones & Company, Inc.

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SCIENTIFIC AMERICAN, INC