Original article

The small hive beetle,

Aethina tumida: a review
WM MICHAEL HOOD
Small hive beetles are now found on three continents and are likely to spread to new regions of the world by means which have yet to be identified.
Since its spread from the African continent, much information has been added to our understanding of this honey bee pest in recent years. Moderately
efficacious methods have been developed to control the small hive beetle in the newly introduced areas, but colony losses are expected to continue
sporadically as beekeepers and scientists work to discover new control alternatives for this pest.
Introduction
The small hive beetle (Aethina tumida) is a
minor honey bee pest in many African
countries. Small hive beetles were first
collected from Old Calabar on the west coast
of Africa and sent to Andrew Murray in
40
London in 1867 for identification. The
beetle is native to sub-Saharan Africa where
they are known as scavengers in honey bee
30,37,55
and received little attention
colonies
prior to their discovery in the USA in 1998.
African honey bee subspecies apparently
possess behavioural traits which allow them
to prevent small hive beetle depredation.

grisella) which are known also for their

function of eliminating or cleaning up dead or
37,55
weakened honey bee colonies.
Damage from small hive beetles is more
apparent in honey bee colonies in the newly
established areas of the world. The beetle
adults and larvae feed on honey, pollen and
30,37,55
Another detrimental effect
bee brood.
left behind by beetles is the spoilage of stored
honey that probably results from beetle
37,55
defecation. The fermented honey left
behind in dead colonies is rejected by honey
bees (which will not consume it) and is
unmarketable by the beekeeper. Beetles are
also recognized for creating pest problems
for beekeepers in and around honey houses
in the new world.

In their native range, the beetles are better

known for their damaging activities around
honey houses and their effect on weak or
stressed honey bee colonies. The small hive
beetle has now become established in new
regions of the world in Australia and the USA
where the pest has become a major problem
for many beekeeping operations. Apparently
European honey bees (Apis mellifera) lack
some of the behavioural traits of African bees
in tolerating the pest and this results in
increased colony losses. Even strong honey
bee colonies have succumbed to the effects of
small hive beetles and died in heavily-infested
areas.

Prior to June 1998, the small hive beetle was

known to occur only on the African
continent (fig. 1). The African countries
reported to have the beetle include: South
37
49
Africa, Botswana, Ethiopia, Kenya, Namibia,
39
52
36
Eritrea, Angola, Central African Republic,
Senegal,45 Ghana,29 Republic of Congo,3
Nigeria,41 Uganda,50 Zimbabwe,55 Guinea61
5
6
Bissau, Congo Republic, Zambia and
57
19
Tanzania, as reported by Ellis.

Prior to 1998, only two significant research

investigations were conducted and published
37,55
on small hive beetles. Their work covered
several aspects of the biology and control of
this hive pest in its native range. Since the
beetle was considered a minor pest of little
economic importance, it was relatively
unknown outside its native range. In Africa
the beetle is often considered less important
than the greater wax moth (Galleria
mellonella) and the lesser wax moth (Achroia

In June 1998, small hive beetles were

collected from honey bee colonies near St
Lucie, Florida, USA, and identified by M C
Thomas of the Florida Department of
20,32,54
Although these were the
Agriculture.
first small hive beetles identified in the
Western hemisphere, earlier unidentified
beetle collections were made in November
1996 and October 1997 in Charleston, South
Carolina, USA. The unidentified beetles were
stored in the Clemson University Insect

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Distribution

FIG. 1. Known distribution of the small

hive beetle in Africa as of October 2003.
Countries where small hive beetles have
been recorded include: 1) South Africa, 2)
Botswana, 3) Zimbabwe, 4) Zambia, 5)
Angola, 6) Tanzania, 7) Democratic
Republic of Congo, 8) Congo Republic, 9)
Uganda, 10) Kenya, 11) Ethiopia, 12)
Eritrea, 13) Central African Republic, 14)
Nigeria, 15) Ghana, 16) Guinea Bissau, 17)
Senegal, 18) Egypt, and 19) Namibia.
Map used with permission of P. Neumann
and was modified from Neumann & Elzen
2003; Ellis 2004.

Bee World 85(3): 5159 (2004) | September 2004 | Bee World | 51

Manitoba Beekeeper's Association and the

local wax rendering plant to contain the pest
so that accidental introductions could be
avoided in the future.9
In October 2002, small hive beetles were
discovered in honey bee colonies in Australia
in Richmond, north-west of Sydney, New
South Wales59 which is also a coastal area (fig.
3). Gillespie et al.28 reported that the beetles
were likely to have been imported into New
South Wales at least six months prior to
their official discovery. An extensive survey
was conducted from October 2002 to
January 2003 in the New South Wales area
and small hive beetles were found mainly
west of the Sidney basin around Richmond
and parts of the lower Blue Mountains.28
Beetle identifications were made from bee
colonies in four regions, including the Sydney
Basin, Cowra, Binalong, and Stroud. Along
with discovery of beetles in managed
colonies,62 13 feral bee colonies in the Sidney
area were confirmed to have small hive
beetles.59

FIG. 2. Known distribution of the small hive beetle in the United States as of December
2003. States where small hive beetles are present and the year of their discovery (in
brackets) include: 1) Florida (1998), 2) South Carolina (1996), 3) Georgia (1998), 4) North
Carolina (1998), 5) New Jersey (1999), 6) Maine (1999), 7) Pennsylvania (1999), 8)
Minnesota (1999), 9) Iowa (1999), 10) Wisconsin (1999), 11) Massachusetts (1999), 12)
Ohio (1999), 13) Michigan (1999), 14) Louisiana (2000), 15) New York (2000), 16) North
Dakota (2000), 17) Tennessee (2000), 18) Indiana (2000), 19) Vermont (2000), 20)
Maryland (2001), 21) Virginia (2001), 22) Delaware (2001), 23) Illinois (2001), 24)
Missouri (2001), 25) Mississippi (2001), 26) Arkansas (2002), 27) Alabama (2002), 28)
Kentucky (2002), 29) West Virginia (2003), and 30) Texas (2003). Map used with
permission of P. Neumann and was modified from Neumann & Elzen 2003; Ellis 2004.
Museum, Department of Entomology,
Clemson, South Carolina, and were later
identified in July 1998 as small hive beetles.
Beetles were collected later in 1998 in many
areas of eastern Florida, coastal South
Carolina and Georgia, and one county in
eastern North Carolina.

fly great distances, perhaps several kilometres,

between apiaries, the widespread discoveries
of the pest in a five-year period in the USA
are thought to be a result of beekeeperassisted movement of beetle-infested
colonies, package bees and empty beekeeping
equipment.

Small hive beetles quickly spread to other

areas in the USA (fig. 2) including Iowa,
Maine, Massachusetts, Michigan, Minnesota,
New Jersey, Ohio, Pennsylvania and
Wisconsin in 1999, Indiana, Louisiana, New
York, North Dakota, Tennessee and Vermont
in 2000, Delaware, Illinois, Maryland,
Mississippi, Missouri and Virginia in 2001,
Alabama, Arkansas and Kentucky in 2002, and
Texas9,35 and West Virginia in 2003.19,39
Although the beetle is thought to be able to

In August 2002, small hive beetles were

discovered in Manitoba, Canada.9 Extensive
surveys were conducted in the Manitoba area
in spring and summer of 2003; no beetle
activity was discovered. The beetles were
unable to establish a population and have not
become a problem in Canada.9 Reports
indicate that the beetles arrived in Manitoba
aboard a shipment of beeswax imported from
MacGregor Wax Works, Hull, Texas, USA.
An agreement was reached with the

52 | September 2004 | Bee World

No conclusive evidence has been reported on

how small hive beetles spread to the USA32
and Australia. The first known small hive
beetle collection recorded in the Western
hemisphere was made by a hobbyist
beekeeper in 1996 in the city of Charleston,
South Carolina, USA, which is home to a
major eastern seaport. Anecdotal reports
from beekeepers in the Savannah, Georgia,
USA, area indicate that small hive beetles
occurred also in their apiaries prior to 1998.
Savannah is another major port city that is
located approximately 135 km (84 miles)
south of Charleston. One suggestion is that
small hive beetles probably arrived in the
Charleston port32 and other ports along the
south eastern USA by human-assisted
movement at approximately the same time
aboard cargo ships loaded with a common
commodity that supported the beetles voyage
from Africa. The likelihood of natural spread
from a single port city in the USA such as
Charleston to become widely spread to
coastal areas of Georgia, Florida and South
Carolina in a two-year period is doubtful.
Beekeepers in the Charleston area are
hobbyist and rarely move bees from the port
city.

Biology
Adult small hive beetles average 5.7 mm in
length and 3.2 mm in width.12 Adult beetles
vary in size which is probably dependent on
food resources and climate.19 Adult female
beetles slightly outnumber and are heavier
than adult males in local populations as
reported by a two-state survey conducted in
the USA.12 Adult small hive beetles are strong
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fliers and are capable of flying several

kilometres59 which aids in their natural
spread. Beetles fly before or after dusk,55 and
males have been reported to fly earlier than
females. They are thought to be attracted to
honey bee colony odours20,60 but may also be
attracted to beetle pheromones which have
not been identified. In olfactometric and
flight-tunnel bioassays, adult small hive beetles
were found to be attracted to volatiles from
adult worker bees, freshly collected pollen,
unripe honey and slumgum.60
Small hive beetles are sexually mature at
about one week following emergence from
the soil.19 Adult females will oviposit directly
on pollen or brood comb if unhindered by
worker bees. Schmolke55 estimated that
female beetles may potentially lay up to 1000
eggs in their lifetime although other estimates
range up to 2000 eggs.59 In a research project
in which nucleus honey bee colonies were
inundated with small hive beetles, female
beetles were observed chewing holes in
capped bee brood and ovipositing eggs on
bee pupae.19 In addition, adult beetles were
reported to oviposit in capped bee brood
through slits they chewed in the side of
adjacent empty cells.19
Small hive beetle eggs are normally laid in
clusters and are pearly white in colour; the
eggs are about 1.4 mm long and 0.26 mm
wide. Female beetles lay eggs in cracks and
crevices around the periphery of the inside of
a highly populated bee colony, but they will
lay eggs in the brood area if unhindered by
adult bees. Most beetle eggs hatch in about
three days but the incubation period can
continue for up to six days.37 Egg hatching
viability is affected by relative humidity.59
Beetle larvae are creamy-white in colour and
emerge from the egg through longitudinal slits
made at the anterior end of the egg.37 The
larval period lasts an average 13.3 days inside
the bee colony and three more days in the
soil. Eischen et al.10 reported beetle larvae
completing maturity in 56 days under
favourable conditions. Beetle larvae are about
1 cm in length when fully grown.37 The length
of mature larvae is variable with smaller
larvae maturing slower and reaching less
length on poorer diets.37 Mature larvae exit
the hive in late evening from 19.00 h to 22.00
h with peak activity at 21.00 h.37 In the honey
house, the relative humidity plays a key role in
beetle larval development,46 so the

FIG. 3. Known distribution of the small hive beetle in Australia as of October 2003. Map
used with permission of P. Neumann and was modified from Neumann & Elzen 2003; Ellis
2004.
manipulation of the moisture level could be
easily integrated into an effective small hive
beetle management programme.
After exiting the colony, mature small hive
beetle larvae enter the soil to pupate where
they reach the pupal stage, a process which
55
lasts about eight days. Female beetles pupate
19
slightly faster than males. Young pupae are
white to brown in colour and are mostly
affected by soil moisture rather than soil
19
type. Soil type was found to have little affect
19
on pupation survivability. Dryer soils impede
pupation success rates. Pupation rates ranged
from 9298% in various soil types provided
19
the soil was moist. This implies that beetle
pest problems can be expected regardless of
soil type in areas where soil moisture remains
high during the year. Therefore, soil moisture
appears to be a major limiting factor in beetle
reproduction thus population buildup. This
may explain partly why small hive beetles are
not a major problem in honey bee colonies in
sub-Saharan Africa because much of Africa
(except equatorial Africa) is semi-arid to
19
arid. The dryer soil conditions would be
expected to have a negative affect on beetle
19
pupation rates. Soil density was found to
affect pupation rates also with high density
soils having a negative affect on pupation

rates. Possible affect of soil temperature on

19
pupation success has not been investigated.
Massive numbers of small hive beetles
perhaps produced from feral bee colonies
have been known to invade and disrupt
63
apparently healthy colonies. Beetle lifespan
and ability to mass reproduce on food
materials found inside honey bee colonies
15
have been investigated. A single mated
female beetle reared on a diet of only pollen
15
produced 591 larvae. Beetle adults survived
37
180188 days when fed honey and pollen,
but only 19 days when fed water and
55
beeswax. Adults feeding on just honey have
15
been reported to survive 176 days but are
not likely to reproduce. Various results were
reported from studies on small hive beetle
lifespan when beetles were deprived of food
55
55
and water; results of two days and 10 days
were reported; seven days lifespan were
reported when adults emerged from the soil
26
and were deprived of food and water. This
suggests that beetles newly emerged from the
soil may live for several days in search of a
new host bee colony or other food source.
Small hive beetles normally overwinter only in
the adult stage and are found within the
honey bee colony cluster where they find
32
food and warmth.

European honey bees (Apis mellifera) lack some of the behavioural traits of African bees in
tolerating the pest and this results in increased colony losses. Even strong honey bee colonies
have succumbed to the effects of small hive beetles and died in heavily-infested areas.
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September 2004 | Bee World | 53

Genetic diversity
Small hive beetle genetic diversity has been
studied in beetles collected from 19962000
in the USA and South Africa.24,25 Two distinct
mitochondrial DNA (mtDNA) haplotypes
were discovered from beetle collections
made in the south eastern USA whereas 13
beetle haplotypes were described from
collections made from southern Africa.
Mitochondrial DNA sequence data from
these separate beetle collections suggest
strongly that a single species exists in both
continents.24 The two beetle haplotypes found
in the USA (NA1 and NA2) are
indistinguishable using the COI gene from
native beetles collected from two countries in
southern Africa: South Africa and
Zimbabwe.24 While these findings suggest
significant similarities between the beetles
collected in the USA and the southern
African countries, conclusive proof does not
exists inferring that the beetles found in the

FIG. 7. Small hive beetle soliciting food

from worker honey bee.
54 | September 2004 | Bee World

FIGS 4-6. Small hive beetle confined in a

propolis prison at the corner of an inner
cover. (4) Bees guarding beetles. (5)
Minimum smoke applied to expose several
beetles. (6) Beetles removed to show
propolis prison.

USA could have originated from other

regions of the African continent.24
The mtDNA analyses conducted by Evans et
24,25
al. yielded an irregular distribution of the
two small hive beetle haplotypes from
collections made over the five year period in
the south eastern USA. The earliest samples
collected in 1996 and 1997 in South Carolina
showed haplotype NA1. Although later
samples from South Carolina in 19982000
included both haplotypes, NA1 continued to
slightly dominate. Samples collected in
Georgia in 19982000 showed a higher
frequency of haplotype NA2, and Florida
samples collected over the same time period
showed roughly equal haplotype frequencies.
Beetle samples collected from South Carolina
and Georgia showed strong biases in mtDNA
haplotype frequencies over the period
19982000 that indicates that the two
haplotypes are not randomly distributed
among beetle populations. Further, results
from the investigations presented competing
hypotheses that beetle haploid frequency
differences indicate two or more separate
introductions into the USA, ongoing selection
at different sites, or genetic bottlenecks exist
as beetles invade new sites.25 Barring strong
bottlenecks as beetles spread to new apiaries
and regions in North America, regional
differences in haplotype frequencies will
erode reflecting the current equal frequency
in Florida.25
No correlation was found in small hive beetle
size and haplotype which indicates that no
selection pressure exists between haplotype
NA1 and NA2 found in the USA.25 Body mass
is often used as a surrogate for competitive
abilities and reproductive success in insects.38

Bee behavioural diversity

Small hive beetle imprisonment behaviour by
honey bees has been well documented in
13,16,17,42,63,65
Guard
recent reports and studies.
bees hold captive adult beetles in propolis
confinement sites and prevent attempted
19
escapes (figs 46). Confinement activity is
likely to play a role in preventing beetle
63
63
matings. Tribe also noted that African bees
are known for their prolific use of propolis to
cover all crevices within the nest to exclude
beetles. European honey bees have been
reported to use almost four-times less
propolis in colonies when compared to Cape
19
honey bees which is consistent with other
1,8,30,53
Cape honey bees were reported
reports.
to hold small hive beetles captive for one to
four days by showing aggressive behaviours
and other sophisticated strategies toward
42
beetles. European honey bees in the USA
have been reported to guard beetles
significantly earlier in age (beginning age 18.55
0.52 days; mean s.e.), to guard beetles
significantly longer (duration 2.36 0.31
days), and to cease guarding beetles
significantly sooner (ending age 19.91 0.57
days) than Cape honey bees in South Africa
(beginning age 20.61 0.38 days; duration
1.43 0.12 days; and ending age 21.04 0.37
16
days). The demonstration of a significant age
difference in relation to guarding behaviour
between the two honey bee subspecies does
not help explain the resulting differential
damage to European and Cape honey bee
16
colonies. Although European honey bees
confine beetles similar to African honey bees,
data suggest this behavioural trait is not as
19
pronounced in European honey bee colonies
which may be the result of less propolis use
in the hive.
Bees have been observed feeding their beetle
captives which is a possible result of
13
behavioural mimicry (fig. 7). Beetles use
their antennae to repeatedly tap the bees'
labium until this behaviour eventually dupes
the house bee into feeding the soliciting
19
hungry beetle.
Imprisoned small hive beetles have been
observed to survive two months without
42
access to food in combs. Beetle survival was
not a result of metabolic reserves because
starved beetles have been reported to die
42
within two weeks. More guard bees were
found to actively patrol beetles at higher
densities and during evening hours than other
17
times of day. Beetle imprisonment activities
by bees are likely to be the first line of
defence against the hive intruders especially at
low beetle densities. Beekeepers may upset
this natural control mechanism when colonies
are opened and manipulated which may lead
to higher beetle activity and stress on
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colonies due to release of beetles from

captivity. The detainment of beetles in hive
cracks and crevices along with the use of
propolis beetle prisons is similar in European
17
and African bee colonies.
When studying European honey bees, Ellis et
17
al. reported finding only 5% of all small hive
beetle adults among combs at lower beetle
densities, but he reported a five-fold increase
in adult beetle activity in comb areas at higher
beetle densities. Higher numbers of beetles
found in comb area at higher beetle densities
may result in increased beetle reproduction
which could help explain the greater damage
19
to European honey bee colonies in the USA.
Natural mechanisms other than confinement
efforts are likely to be responsible for making
African bees more tolerant of small hive
19
beetle activities. African bees exhibit more
aggressive behaviour against free-roaming
23
beetles. The hygienic behaviour of removal
of beetle eggs from capped brood cells may
be an important resistance mechanism by
19
honey bee colonies. African bee colonies
readily remove exposed beetle eggs and
44
larvae which may be an important trait in
the genetic control of this hive pest. But
investigations have shown that both Cape and
European honey bees removed beetle-egg
infected capped brood at similar rates in
19
controlled studies. A practical assay for
testing colonies for removal of brood cells
infected with beetle eggs has been
19
developed. The stimulus used by bees to
identify beetle eggs under capped brood cells
19
remains unclear.
Honey bees in South Africa are well known
for their absconding tendency whenever small
30
hive beetle numbers increase in a colony,
whereas this trait was thought to be absent in
European honey bees. However, results of
19
recent investigations reported high
absconding rates (6 out of 10 colonies) in
European honey bee colonies when artificially
challenged with large numbers (100 adult
beetles added per day) of beetles over a 14day period. In experiments, all 10 treated
colonies, including those absconding and not
absconding, uncapped and discarded most or
19
all capped pre-pupae and pupae. These
results indicate that European honey bees do
respond to large populations of beetles by
having a high, prepared absconding rate
19
similar to African honey bee subspecies.
However, a slower natural increase of small
hive beetles in European honey bee colonies
might yield different results.

Economic importance
The small hive beetle is considered to be of
little economic importance in its native range
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Beetle imprisonment activities by bees are likely to be the first

line of defence against the hive intruders especially at low
beetle densities. Beekeepers may upset this natural control
mechanism when colonies are opened and manipulated which
may lead to higher beetle activity and stress on colonies due to
release of beetles from captivity.
of southern Africa where it is listed as a
threat only to weakened or stressed
37,55
colonies. African bees are capable of
preventing the beetle from breeding in the
hive as long as colonies remain strong.56 In
contrast, beetle infestations in the south
eastern USA have affected even robust bee
colonies which required control measures by
the beekeeper.20 Beetles feed on honey, pollen
and brood in bee colonies and have been
implicated often in both colony mortality and
increased absconding rates.24 A quarantine on
movement of honey bee colonies was
established in Florida, USA in June 1998, but
it was soon withdrawn by the Florida
Department of Agriculture and Consumer
Services.27
The estimated losses to small hive beetles
experienced by beekeepers in the USA in
1998 were US$3 million.23 Losses were in the
form of colony destruction and damage to
stored honey supers in honey houses. Some
commercial beekeepers in the USA reported
losing thousands of bee colonies and
associated equipment to beetles the first few
years following their discovery.59,65 Mixed
reports are coming from Australia as to the
level of damage the small hive beetle is
causing in that country,62 although strategies
for prevention and management of the beetle
have been developed and provided to the
beekeeping industry.28 Reports indicate that
beetles have not caused significant damage in
Australia66 when compared to damage caused
in the USA, especially coastal south eastern
USA.32 A beetle survey conducted in managed
honey bee colonies from October 2002 to
January 2003 in the New South Wales area of
Australia reported 120 positive detections
out of more than 1000 samples received.28
Seven drought years in the Australian beetleinfested areas restricted movement of
colonies which may have resulted in slowing
the spread of the beetle.58 The major losses
have been a result of the negative effect on
overseas and domestic package and queen
bee markets. White66 reported that stressed
bee colonies suffering from European
foulbrood are prone to result in major small
hive beetle problems.

The queen and package bee production

business has been negatively affected by small
hive beetles in the USA also. Beekeepers have
concerns over beetles spread in queen cages
and packages.4,19 Concern over spread of small
hive beetles to the UK has been reported.4
Favourable conditions required for beetle
survival are met in many areas of the UK.
Therefore, the risk management
recommendations for small hive beetles in
the UK include prohibition of bee imports
from infested countries.4
Concern over possible small hive beetle
damage to other commodities such as fruits
has been raised. Studies have investigated
beetle reproduction on alternate food
sources such as fruits.10,19 Beetles regenerated
when offered a diet of avocado, cantaloupe or
grapefruit in confinement.10 Laboratory reared
beetle adults were fed rotten and fresh kei
apples (Dovyalis caffra) and survived an
average 58.6 days and 63.9 days, respectively.15
Average number of offspring produced from
three mating beetle pairs after feeding on
rotten kei apples in laboratory tests were
significantly less than the average number of
offspring produced from three mating beetle
pairs fed on pollen comb (10.6 vs. 1096.4).15
The poor reproductive success of beetles
feeding on fruits is likely to be a result of
minimum nutritional requirements being met,
but there is the possibility of beetle
regeneration on fruit in the wild when no bee
colonies are present.15 No record exists
which reports successful beetle regeneration
on fruits or vegetables in field conditions.
Since small hive beetles can survive for
several days on various fruits,10,15 there exists
a strong possibility that beetles can be
transported by fruit truck or cargo shipments
to non-infested regions of the world.
Bumble bee and other non-Apis species are
additional concerns as possible threats to
small hive beetle invasion.2 In controlled
laboratory studies, beetles regenerated on
colonies of bumble bees.2 These investigations
were conducted in confinement and no one
has reported finding small hive beetles in
natural bumble bee colonies, but surveys have
not been conducted to refute this possibility.
When beekeepers move beetle-infested
September 2004 | Bee World | 55

honey bee colonies from location to location

for commercial pollination purposes, they
may leave behind great quantities of beetle
pupae in the soil which emerge to seek and
find a suitable food source. Fortunately, it
appears that small hive beetles are host
specific to honey bee colonies and if the
emerging beetles do not find a new host
colony soon, they are likely to perish. There
is, however, the possibility that small hive
beetles may be attracted to ground-nesting
bumble bees because of similar odours (bee
brood and honey) as honey bee colonies. This
could prove detrimental to bumble bee
colonies during the warmer seasons of the
year. Although bumble bees do not
overwinter as colonies in many regions of
world, the beetles presumably would perish
along with the colonies for lack of food and
warmth.

FIG. 8. Battery operated vacuum for small

hive beetle removal (Hausherr's Machine
Works, 1186 Old Freehold Road, Toms
River, New Jersey 08753, USA).

Biological control
Biological factors, such as natural pathogens,
may play an important role in small hive
beetle control, especially in the beetles'
endemic range of sub-Saharan Africa.
Scientists have reported unknown fungal
pathogens possibly causing beetle mortality in
19,37
laboratory studies. Recent investigations
have been conducted in laboratory trials to
identify possible fungal pathogens that are
responsible for beetle mortality.19 Several
fungal pathogens were identified from dead
beetle larvae including soil dwelling fungi,
Aspergillus flavus and A. niger, which are known
for attacking other soil infesting insects.
Three other saprotrophic fungi (Clonostachys
rosea, Gliocladium catenulatum and Mucor
plumbeus) were identified from the surface of
dead small hive beetle larvae in experimental
studies. Further work is needed in this area
of biological control to identify those
pathogens that may be responsible for beetle
death, especially in the beetles' endemic range
of southern Africa. This biological control
mechanism if developed has potential
application in newly introduced areas of the
19
world.
Additional biological agents may play a role in
controlling small hive beetles in some areas.
Some potential agents are soil infesting
nematodes, parasitic wasps and flies, and
predators such as ants. The imported fire ant,
Solenopsis invicta, infests much of the current
beetle-infested range in south eastern USA.
The imported fire ant has been observed
feeding on mature small hive beetle larvae as
31
they enter the soil to pupate. Fire ants may
reduce beetle activity in some areas but little
is known about this predatorprey
relationship.
56 | September 2004 | Bee World

FIG. 9. PVC pipe upper hive entrance.

FIG. 10. Plastic box trap for small hive

beetles mounted on hive body frame.

present is recommended to kill all life stages

of the pest. The application of sugar patties to
bee colonies for antibiotic or non-antibiotic
purposes may lead to small hive beetle
65
65
problems. Westervelt et al. reported
increased beetle incidence in colonies having
sugar patties. In beetle-infested areas,
beekeepers should be careful when feeding
colonies sugar water or corn syrup inside
hives. Beetle larvae have been discovered in
great numbers in sugar water placed inside
boardman feeders where beetles are well
protected from bee aggression. Push-in
screened queen introduction cages are not
recommended for use in heavily beetleinfested areas. Beetle adults and larvae enter
the screen cage and are well protected with
only the queen present.
Good sanitation is recommended around
honey houses to prevent small hive beetle
32,37
damage to stored comb. Beekeepers
should remove wax cappings, other wax
materials and equipment containing bee
pollen. Pollen traps should not be left on
colonies over extended periods of time
because the unprotected pollen will provide
the beetles with needed protein for
15
regeneration. In the honey house, honey
should be extracted from supers within 23
days to prevent beetle damage. Maintaining a
relative humidity of 50% or less in honey
59
houses will result in beetle egg dessication.
Selecting apiary sites that have drier soil
conditions is recommended for small hive
beetle control; open, sunny sites are
recommended. In commercial agricultural
settings where fields are often irrigated,
beekeepers should place bee colonies several
meters from irrigated areas to minimize
beetle regeneration because moist soil
19
conditions promote regeneration.

Chemical control
Cultural control
There are many cultural practices that
beekeepers may use to minimize small hive
beetle problems. Reducing colony stress
conditions and maintaining strong productive
colonies are highly recommended, especially
in areas where beetles are problematic. Any
practice which helps to maintain wellpopulated honey bee colonies that reduces
the comb-to-bee ratio37,55 and excludes
beetles from the brood area is
recommended. Good bee management
practices that reduce the likelihood of brood
disease, mite problems, wax moth activity,
failing queens, excessive swarming, oversupering and colony starvation are
recommended.31,32 Freezing dead or weakened
colonies and empty supers that have beetles

Chemical products for small hive beetle

control have been developed but varying
37
results have been reported. Lundie reported
the use of carbon disulfide as a fumigant to
control beetles in stored comb.
Paradichlorobenzene has been suggested also
as a fumigant for beetle control in stored
39
comb. Household bleach was recommended
as an effective material for killing beetle adults
46
and larvae in honey houses.
Various soil treatment materials have been
tested to control small hive beetles when
they enter the soil to pupate. In South Africa,
soil treatment tests were conducted using
HCH (benzene hexachloride), carbaryl,
55
chlordasol and salt solutions. HCH and
chlordasol were found to be more effective in
the study. Gard Star (a.i. 40% permethrin, Ywww.ibra.org.uk

Tex Corp, Cody, Wyoming, 82414, USA) has

been registered in many beetle infested states
in the USA since 1999. The product is a soil
drench material that when mixed with water
and applied to the soil in front of bee
colonies will kill beetle larvae and pupae.
Gard Star has also been recommended to
treat old apiary sites where beetle infested
colonies have been removed and beetles have
been left behind to emerge from the soil.7 In
Australia, the National Registration Authority
(NRA) issued a permit to allow soil
treatment with Farmoz Permex EC
insecticide plus other registered products
containing 500g/l permethrin.66 Pettis &
Shimanuki47 recommended pesticide soil
treatments under and extending out 90180
cm from the hive in all directions to control
beetles. Placement of colonies on stands or
blocks is recommended to prevent soil
pesticide fumes from entering hive
entrances.32
An in-hive small hive beetle control product,
Check Mite + (a.i. 10% coumaphos plastic
strip, Bayer Corp., Shawnee Mission, Kansas

66201, USA) has been available in the USA

since 1999. A single strip of the product cut
in half should be attached underneath a piece
of 101.6 101.6 mm (4 4 inches)
cardboard which the backing has been
removed to expose the corrugations. Beetles
are attracted to the dark, corrugated area
underneath the cardboard that is placed to
the rear of the hive at the centre of the
bottom board. Beetles come in contact with
the pesticide while seeking refuge and remain
there long enough to receive a lethal dose.
The product is effective when beetles are
active but is ineffective during cool weather
when beetles confine their activities in or
near the bee cluster. The product is labelled
for use only when bees are not making
surplus honey and honey supers must be
removed during treatment.19,32
Neumann & Elzen43 suggested the idea of
'natural' chemical control of small hive beetles
by the use of pheromones. Male-produced
aggregation pheromones have been
described48 and used for control of other
beetle species in the family Nitidulidae.

Studies have reported that male small hive

beetles enter honey bee colonies before
females19,22 which suggests the existence of
similar aggregation pheromones that may play
a role in colony invasions.19 Laboratory
research is being conducted to identify the
chemical attractant air-borne volatiles from
various sources inside honey bee colonies to
develop lures and traps for small hive beetle
control in the field.60 The development of
colony treatment thresholds for small hive
beetle management is needed greatly.19 At
present, no guidelines exist for beekeepers to
determine the beetle infestation level inside
bee colonies that warrants treatment. The
development of beetle treatment thresholds
will enhance the integrated management of
this pest and result in fewer treatments and
thereby less cost to the beekeeper and less
chance of hive product contamination.

Physical and mechanical control

Several physical methods are available to
beekeepers for controlling small hive beetles
although some are time consuming and
efficiency is questionable. Removal of beetles
from bee colonies by vacuum (fig. 8) or by
hand is possible but these methods are very
time consuming and repeat visits to the same
colonies are sometimes necessary, especially
when beetle infestations are high. Anecdotal
reports from some hobbyist beekeepers
claim excellent beetle control from smashing
adult beetles with a hive tool, but the process
takes several minutes per colony per visit,
which makes this method unrealistic for most
beekeepers.
When honey bee colonies die with significant
numbers of beetles present, the hive
equipment left behind is often covered with
slimy, fermented honey and live beetle larvae.
Investigations46 have been conducted to
determine effective measures the beekeeper
may take to salvage the equipment. Several
household products including white vinegar,
regular bleach, dishwashing detergent, and
vegetable oil have been tested to kill beetle
larvae and test for honey bee acceptance of
the affected comb. Bleach (50%) was the
most effective and fastest acting material
killing 100% of the larvae within four hours.
Dishwashing detergent (1%) killed 85% of the
beetle larvae within 24 hours of treatment.

FIG. 11. West Beetle Trap (Dadant & Sons, Inc, 5 South 2nd Street, Hamilton, IL 623411397).
www.ibra.org.uk

Moving bee colonies frequently to new apiary

sites has been reported to reduce beetle
problems32 but this is extremely labour
intensive. Selecting apiary sites that are
exposed to full sunlight has been suggested
for beetle control. This may be directly
related to the theory that less soil moisture
greatly reduces beetle survivability.19
September 2004 | Bee World | 57

Small hive beetle trapping investigations have

21,34,51,55,60
Attempts have been
been reported.
made to exclude beetles from entering or
55
exiting bee hives. Smolke conducted
preliminary trapping investigations with three
simple in-hive beetle traps. Two of his traps
were designed to be placed above the inner
cover and the other was placed at the rear of
the colony on the bottom board. His efforts
were unsuccessful although his experiments
were hampered by cold weather when
beetles were inactive.
21
Elzen et al. developed a 7.6 litre (2 US
gallon) plastic bucket trap for adult small hive
beetles. The trap was constructed of four
pieces of 8-mesh hardware cloth each glued
to a 7-cm (diameter) hole cut in the side of
each quadrant of the bucket. The opening in
the screen was large enough for beetle
passage, but too small for bee passage. The
traps were baited with various combinations
of hive products and bees including honey,
pollen, excised pieces of comb honey, bee
brood and live adult bees. The baited hives
were positioned in beetle-infested apiaries
and checked at 24-h and 48-h periods. The
combination of honey, pollen and live adult
bees was found to be the most attractive to
21
adult beetles. However, these traps proved
to be ineffective beetle control devices in
apiaries most likely due to competing hive
odours emanating from nearby honey bee
colonies.

The use of an upper hive entrance to control

14,18
small hive beetles has been investigated.
Hood & Taber33 first suggested the idea of
using an upper hive entrance to control this
14,18
pest. Ellis et al. conducted trials to test 1.9
and 3.8 cm (interior diameter) polyvinyl
chloride (PVC) pipe upper hive entrances
while the lower hive entrance was closed (fig.
9). Inconsistent beetle control was reported
with the upper hive entrance and fewer adult
bees and brood were produced when
compared to the normal hive entrance
colonies. Other problems associated with the
upper hive entrance when using a wooden
bottom were impaired thermoregulation,
excessive floor debris and poor water
14
drainage. Some of the negative effects were
mitigated by using a modified screen bottom
board but further research was suggested to
investigate the use of an upper hive entrance
18
for small hive beetle control.
Hood & Miller34 tested an in-hive plastic box
trap (152 80 25 mm) containing a
removable lid which had small vents to allow
beetle entry (fig. 10). The trap was designed
to be fastened with screws to the bottom bar
of a hive frame. Various liquids were placed in
the trap to investigate beetle attractiveness
and lethality. Alcohol (95% ETOH and 5%
58 | September 2004 | Bee World

water), beer, ethylene glycol, mineral oil,

honey and cider vinegar were tested. Beer
proved to be a good attractant in field colony
tests but crusted over during warm weather
and lethality was poor in laboratory tests.
Cider vinegar showed good attractiveness in
field colony tests but lethality was poor in
laboratory tests. In contrast, mineral oil
proved to be highly lethal in laboratory tests
but showed little attractiveness in field colony
tests. Although no materials tested provided
adequate beetle control, some materials
proved to be good attractants but
unfortunately the trap was not designed to
prevent beetle escape. Further development
of a one-way beetle trap that is convenient
and efficient may provide a safe and
inexpensive method for beekeepers to
manage small hive beetles. Other traps have
been proposed for small hive beetle control
such as a wooden box (76 76 254 mm,
filled with sawdust soaked with food grade
mineral oil) suspended from the landing board
to trap mature beetle larvae as they exit the
hive to pupate in the soil.51 Dadant & Sons,
Inc. (51 South 2nd Street, Hamilton, IL
62341-1397, USA) currently markets the
'West Beetle Trap' (fig. 11) that is a plastic
tray which when one-fourth-filled with
vegetable oil is placed directly on top of the
bottom board of a hive. A spacer is included
with the trap that is used to raise the hive
body 2 cm to give extra clearance for the
tray. A slotted cover (296 openings, 2 45
mm each) fits tightly on the tray that allows
beetle entry but excludes bees. All supers
must be removed from the bottom board to
install the trap, and the hive must be level to
prevent the oil from leaking from the tray and
killing bees.64 An additional benefit of the
West Beetle Trap is that varroa (Varroa
destructor) fall from the colony into the oil and
perish along with the beetles,64 which is an
innovative approach to controlling a two pest
complex.

Conclusions
Small hive beetles are now well established in
new regions of the world. The likelihood of
further spread to additional territories and
continents where Apis species are endemic is
expected. This pest may become a significant
problem in countries where conditions are
favourable for beetle regeneration. Small hive
beetles have the potential to survive in any
region of the world excluding those having
extreme ecosystems such as deserts or polar
areas.19
Effective small hive beetle management
recommendations will include good
beekeeping practices that promote and
maintain healthy and well-populated honey

bee colonies. There will be occasional

outbreaks of beetles that will affect even
strong honey bee colonies when conditions
are favourable for beetle reproduction. Small
hive beetle fecundity may be tied directly to
local environmental conditions such as rainfall,
19
temperature, soil moisture and soil density.
Beetle problems may be exacerbated by
other honey bee maladies such as disease and
parasitic mites which may result in a
synergistic effect on colonies. Continued
efforts to develop effective and safe methods
for small hive beetle management will offer
the beekeeper several options to control this
hive pest.

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WILLIAM MICHAEL HOOD

Department of Entomology, Box 340315, 305 Long Hall, Clemson University, Clemson, South Carolina, USA
mhood@clemson.edu
www.ibra.org.uk

September 2004 | Bee World | 59