Download Host selection or mate selection?

Proceedings of the 7th Internatumol
Workmg Conference on Stored-proiuct Protectwn - Volume 2
Host selection or mate selection? Lessons from Prostephanus
truncatus, a pest poorly adapted to stored products
Hodges R
J1
,BIrkmshaw L. A. land Smrth R H.2
Abetract
Prostephanus truncatus (Horn) ISpnmanly a wood bonng
msect that IS pre-adapted to mfest only certaun stored
products, specifically maize and dned cassava. LIttle IS
known about host selection of ItS typical woody hosts
However, a number of studies of ItS response to food
matenals, pheromone biology and sexual interactions have
shed light on how stored-product hosts are colonized
Pnmary host selection ISachieved by either males or females
at random usmg the 'land and choose' behaviour while
secondary selection occurs as either sex IS attracted to the
pheromone produced by males that have found food Females
and males are preferentially attracted to the pheromone
released by particular males although on contact females
select mates accordmg to other features For females, host
location would appear to motivate response to the pheromone
although mate selection could be an additional advantage of
bemg able to distmguish between the pheromone secretions
of different males.
Aggregation pheromones are a common feature of many of
the Coleoptera found mfestmg stored products. Compared
WIth other species, P truncatus IS relatively poorly
adapted to the 'explOItationof stored products, and may there
offer some insights mto the early stages of the evolution of a
well-adapted storage pest.
Introduction
Prostephanus
truncatus
(Horn)
(Coleoptera:
Bostnchidae) ISa well-known pest m farm storage although
generally restncted to only maize and cassava. It was
mtroduced into Mrica from Meso-Amenca m the late 1970s
and smce then has spread WIdelym that contment (Hodges,
1994) where It has been the cause of substantial food losses
P. truncatus appears to dIffer from many of the well
known storage pests III that It has large populations away
I Food Secunty Department, Natural Resources InstItute, Ulliversity
of GreenwIch, Chatham Mantlme, Kent ME4 4TB, UK
2
Department
of
BIologIcal Sciences,
Ulliversity
Ulliversity Road, Leicester LEI RH7, UK
of
Leicester,
from storage systems where It IS believed to live on woody
hosts Observations m both Kenya'( Nang'ayo et al , 1993)
and MeXICO(Ranurez-Martmez et al , 1994) confirm that
this pest can reproduce on dead wood, and m a laboratory
study Nang'ayo (1996) demonstrated vanable abrhty to
breed on the wood of 24 species of tree, belongmg to eight
famihes, at moisture contents ranging from 8.5 % to
14.8% However, desprte considerable efforts It has rarely
been found m the wild, even m places where high catches in
pheromone traps indicate that the beetle IS abundant. In
both MeXICOand Kenya It has been suggested that common
hosts of P trumcatus include branches of trees that have
fallen victim to girdhng cerambycids (Ramrrez-Martmez et
a1. , 1994; Nang'ayo , 1996)
Evidence that P.
truncatus
Is Pooryly Adapted to
the Infestation of Stored Products
The strong aSSOCIatIOn
of P truncatus With woody hosts
has led biologists to conclude that it has evolved as a wood
borer (Chittenden, 1911; Nang'ayo et a1. , 1993; RamirezMartmez et al , 1994). It IS known that many typical
storage pests, includmg the closely related species
Rhyzopertha dommica , are strongly attracted from
considerable distances to the odours of stored products
(Barrer , 1983), a behaviour WhIChthey do not share with
P truncatus (Hodges, 1994) Prostephanus truncatus
would, therefore, appear to be a forest species that in some
measure IS preadapted to life in maize and dned cassava
should It reach them by chance Its explortation of stored
products IS sporadic and concentrated within relatively few
stores where It can cause enormous damage (Hodges, 1986
and 1994; Markham et aI., 1991) compared WIth welladapted storage pests such as Siiophalu« zeamais ,
Tribolrum
cosianeum.,
Ephestia
cautella
and
Rhyzopertha domuHca, WhIChpose a constant threat of
mfestatIon and whIch may attack a Wide range of food lots
wlthm a gIven localIty
A tentative host selection hypotheSIS for P truncatus
has been proposed by Hodges (1994).
An Important
component of this hypotheSISIS the aggregation pheromone
released by the male. There have been some recent studIes
on the sexual behaVIOurof P truncatus (BIrkmshaw,
1998) and new observations on the pheromone mteractIons
of thIS pest (SmIth et al , 1996; Scholz et aI., 1997 a&b;
1788
Proceedings of the 7 th Internatwnal
Hodges et al , 1998; Hodges and Dobson,
therefore
m press).
timely to update the host selection
to
because unrelated
mdrvtduals (competitors)
may have m the
fitness mcreased.
A mutation
at the host and/or
m
mate selection by females.
Selection thinking
of
thmkmg when discussmg
features
such
and
the
the
mutation
would
as
aggregation
population
or species.
along the lines of 'aggregation
pheromone
more
frequently
through
Whatever
group-selected
are untenable for one
traits are not evolutionanly
stable and Will be rapidly replaced
by mutant
benefit
the
individual
rather
than
Illustrate tlus general pnnciple
functions of aggregation
traits
group.
by reference
unrelated
mdividuals,
e.
1.
km
the food medium through communal feedmg (Wood 1982),
(unphcitly
suggesting group selection)
than
or ma "condinonmg ' to Improve the quality of
selection),
or ina attraction
the
spread
mdrvidual SIgnaller. ThIS benefit might be, for example, via
inclusrve fitness (If close relatives were attracted to food
has evolved to help other members of the species fmd food'
simple reason:
mevitably
mamtamed m the face of mutation If there IS a benefit to the
that the umt of selection IS the individual
msect or gene and not the group,
Loose arguments
mcrease the fitness of
the mutant relative to other members of the population,
It IS therefore clear that pheromone SIgnalling can only be
the
We Wish here to emphasize the central tenet of
modern evolution:
will have thetr
causmg an individual not to
produce pheromone would, therefore,
have not always followed the
logic of modern evolutionary
function
The
population.
Stored product entomologists
pheromones.
by a kairomonal effect).
at the same time,
and,
attraction
adaptive
because a predator IS attracted
SIgnaller will certamly suffer a reduction in relative fitness
to
consider what role pheromone
of beetle aggregations
It IS
hypothesis
mclude the latest observations
development
Working Conference on Stored-product Protectum - Volume 2
ultimate
that
of a potential
the proximate
benefit
mate (Birkinshaw
mechamsm,
m terms
there
of mcreased
1998)
must
mdividual
ThIS lOgICIS known as "selection thmkmg ' (Charnov,
We WIll
be an
fitness.
1982)
and has been expressed very lucidly and forcibly by Dawkms
to the possible
(1976)
pheromone.
Primary Host Selection
Beetlesdisperse by
/ftYlng
Or",alkJn~
In P.
Pheromonedetected
Pheromonenot detected
!
especially
Beetleselects substrate
for bonng
!
Male
Female
locat! food
locat! food
Mall arnve
not
'Tted
walk or fly from food sources,
are depleted
1997).
Most flight actrvity
around
dusk
(Tigar
of the beetle
et al.,
1993),
personal
observation ) and most
already
mated
are
IS therefore
dispersal
Female
P.
(Scholz,
truncatus
(maize or cassava) by Prostephanus truncatus
attracted
food source,
that all mdividuais
in a population
produce
when they fmd food. ThIS pheromone
mdividuais
but
also proVldes
for entomologIsts
of both sexes to the food,
a
attracts
male and female beetles alight on a surface and make test
soap,
sIgnaller)
The
producing
SIgnaller
absolute terms (perhaps
may suffer
the
pheromone
reduced
fItness
(a
m
because energy used to syntheSIse
pheromone IS not available for growth and reproduction,
leather,
penetrate
nature
or
1).
The test
burrows are usually not more than about 1 cm m depth and
may be made mto almost any solid matenal,
mdividual
et
For primary host selection,
burrows m search of SUItable food (FIgure
and thIS
mvestIgating
to stored products over a long range (Wright
benefIt
an
an
truncatus was that the adult beetles are not specifIcally
somehow benefIts other members of the speCIes but IS of no
to
It
1998).
mtended for
for the selectlOn of new mates
al , 1993; Tigar et al , 1994).
unrelated
(Birkmshaw,
hkely that such flight is pnmanly
to a new
opportunity
P.
Suppose
S. Addo
have an extended
Reproduction
Strategy for the selection of stored products hosts
pheromone
dispersing
1997;
penod and have been found to mate multiply
One of the fIrst surpnses
Fig. 1.
IS also
Such flight
(Birkinshaw
reproductive
)
Males stop pheromone
productIon
there
1991).
females
unpublished)
contmres
Femalesamve -
population
IS concentrated
but
a peak at dawn (NovlIlo,
WIth the same and different partners
~~~~~~~e
or msect
1986; Fadamiro , 1995a; Scholz,
smce matmg can occur m the tummels of the food source
Backsout
IncreaseIn populatIon
densrty decreaseIn
foodquahty
Femalesarnve
these
would not appear to be required for the act of mating itself
\'oo~
Beetlesarnve at
7rornone ~ce
when
sometimes
Food
Eggs laid
Pheromrne released
dispersal away from stored product hosts
density IS high (Hodges,
Beetlemakestest burrow
In searct of food ~
/
truncatus,
occurs as adult P. truncatus
of
etc.,
that
Short-range
(1997b)
test-bormg
behavlOur
chemIcal/visual
1789
plastic,
IS not
but
the
suggests
cues are not a prereqUIsIte
such as wood,
too hard
to
mdIscnminate
that
speCIfIC
to tunnellmg
Proceeimq« of the 7th Internatwnal
Working Conference on Stored-product Protection - Volurne 2
If a beetle falls to find food in Its test burrow then
behavior.
It will back out and continue the search.
It IS not certain
which food matenals
truncatus
P
most m search of. It can digest cellulose (Vazquez-Ansta
al. , 1997),
et al , 1994;
al , 1998).
Similarly,
not volatile and IS normally
starch.
In expenmental
& Dobson,
grain ( Hodges
male P.
truncatus
truncatus,
although the median
number of beetles attracted
dunng this experiment
two (Birkmshaw , 1998)
Interestingly,
these beetles was significantly
m favour of females (64%).
one week,
truncatus
possibility
m fhght
(Fadamlro
that the volatiles
et
ai,
emanatmg
product insects might also attract
P
studied
by Scholz et al , (1997b)
results.
However,
these researchers
beetles,
1998)
The
from other
stored
truncaius
wtth
walking beetles respectively,
their
76%
m
l'fevlce
traps.
smce P.
sex ratio
1S beheved to favour the hosts that cerambyc1ds
have attacked
of pheromone
favour
by wh1ch
behaVIour
may
and
be
concentration
thereby
poSSIble that males,
the small colony
male P
attract
Unhke
they release
conventional
perpetuatmg
truncatus
only t!J.e oPpos1te sex
aggregation
(Borden,
pheromones
are
an aggregahon
to that
of the
attractIve
to
1985) and m the case of P.
are
medIated
from
the
m flight
However,
sex ratio dIstortIOn m
by
that
a
although attracted
m
traps.
on sex ratio dIstortIon
In
landmg
pheromone
It IS therefore
to the pheromone,
view
and hence
of
similar
of the catch m creVice
where the beetles would locate
the trap by walkmg rather than flymg,
sexes
male
lower
dIstance from It than females,
traps (Hodges et al , 1998),
SIgnaller,
truncatus,
beetles
the sexes
so suggested
under-represented
observatIon
whIch
both
of
threshold than that for females.
settle at a greater
so as soon
sex pheromones,
Samples
lure showed greater
of females
tIme some of her progeny would have reached adulthood and
pheromone.
of the catch
65% m fhght traps and
Fadanuro (1995) reported no differences
could mate with her and each other,
as they locate a food source
the preponderance
Scholz( 1997) found that traps loaded With greater quantitIes
by herself as she would be albe to
On the other hand,
In flight traps or crevice
designed to catch flying or
actIvIty or duratIon of fhght between
Once pnmary host selectIon has taken place, a female can
can not create a colony in the absence of females,
60
female biased
population being trapped WIth the crevice traps had an equal
truncatus
lay fertIhzed eggs for up to 46 days (L1, 1988),
pheromone,
and left for
of about
whose sex ratio was also sigmficantly
was female (Hodges et al. , 1998);
studies to typical forest pests such as girdling cerambycids,
explOIt the food reserve
population
(65 %) (Scholz et al , 1997b).
has been
wh1ch mIght well prov1de more posItive results
an average
traps WIth synthetic
no conclusive
plan to extend
accrued
was only
the sex ratio of
of suitable food. It is probably for tlus reason
of P.
a single
feeding on marze has been observed to
Maize cobs imtially baited With a single male,
that maize
of the
beetles to fly. only when in
towards the pheromone.
attract up to 35 other P
volatiles.
the orientation
1998).
Fadarruro ( 1995) found that the presence
These volatilize would thus be no sure SIgn of the presence
volatiles do not appear to influence
to males
In a penod of one night m the field (Ghana),
Starch IS
encountered
maize
1993 a&b) .
fhght do beetles orientate
et
hidden below layers of plant
that would errut commonly
inside
by (PIke,
pheromone will not stimulate
when maize and dned cassava roots are
the latter being almost enttrely
matter
once
Starch can support the high rates of population
increase observed
infested,
Borgemeister
even virgm females would not respond
IS
1990; Ramirez-
Nang'ayo , 1996;
close
studies,
et
but It would seem that starch IS a particularly
Important component of the diet (Detmers,
Martmez
released
thIS effect may not
be confmed solely to landmg behaVIour
mitIate
Successful
the process of secondary host selectIOn.
host selectIOn
fmdmg food and attractmg
by males
depends
eIther
a mate or on respondmg
on
to the
SIgnal of another male. In the latter case, they may have to
Secondary Host Selection
compete for both females and food
For males the lowest
nsk may be to respond to another male smce thIS WIll enable
The aggregatIon
pheromone released by male P.
IS a blend of two components called Trunc-call
trU'ncatus
1 and Trunc-
call 2 (Cork et al , 1991; Dendy et al. , 1989,1991).
release
their
pheromone
from wlthm
the host,
a rapid locatIOn of a food source
chances of attractmg
Males
a female
and may Improve
smce
there
the
WIll now be
multIple pheromone SIgnals from one locatIon. In practIce,
and thIS
dIffuses to the extenor
through the small holes made by the
SIgnals, but remam far enough away to aVOId competItIon
beetles's
Pheromone
WIth other males for any female that is attracted
tunnelling
males on malze gram IS vanable,
collected
amountmg
from
sohtary
to a dally total
components
rangmg
for 1.0fJ-g
conSIstent dIfferences
between
males m both amount
for both
ratIo of components
(Hodges and Farman,
Whether
to 4. OfJ-g wIth
unpubhshed)
to hIm
or not males actually settle at a speCifIc optimal
distance from each other could be easIly tested by further
and
It
expenmental
work or fIeld observatIon.
WhIle males appear
to keep theIr dIstance from other males, females also appear
would appear those only beetles m the process of dIspersal
to aVOIdor repel each other (Hodges and Dobson, m press):
respond to pheromone
perhaps m both cases the behaVIour IS to hmlt same-sex
smce,
once mSIde a host,
males nor females WIll leave It m response
neIther
to pheromone
a
male needs to be close enough to benefIt from the combmed
competItIon.
1790
Although
males
respond
to the pheromone
Proceedings of the 7th International
signal of other males, the opportumty
be bnef as, once a female arnves
Workmg Conference an Stored-product Protection - Volume 2
for them to do so may
m close proxmuty
of females,
to a
could be adaptations
to hmit loss of patermty
from other male suitors and to reduce the nsk of predation.
male, pheromone production declmes rapidly (Smith et al. ,
The latter IS a particular
1996; Scholz et al , 1997a; Hodges and Dobson, m press)
IS also
This behavior
Teretrius nutreecene (Lewis }, formerly Teretrioeoma
niqrescens (Rees et al , 1990; Boeye et al , 1992).
volatile:
IS mediated
males
amputation
The
by a female factor that IS non-
will respond
to It even
after
antennal
(Smith et al , 1996)
fmdmgs
pheromone,
that
only
In summary,
males
produce
for
the
host selection
predatory
in the
a dechrnng
food source
and that males cease to SIgnal m the
Thereafter,
there may be pnmary selection,
presence
females,
the response to as yet umdenufied
of
adult
ornaments,
be they
visual,
m the relative
examples
reproductive
of
such
elaborate peacock's
truncaius
or
traits
pheromone
signalling
and cuurrent
FIeld tnals have demonstrated
attracted
particular pheromone
truncatus
however,
these
charactenstics
results
from the
to the pheromone
pheromone
come mto contact with a
female, pheromone production IS stopped;
SImilar behaviour
only females,
If females
of potential
females
for their
by males
while males and females
encounter
the aggregation
pheromone
IS possible.
are normally
However,
already
mated
screemng
as dispersing
and as even
SUItable food source,
It seems
likely
that
motivates the response to the pheromone.
some P
matmgs
mates
virgin
females WIll not respond to the pheromon once they are in a
and Farman,
that
more
It
1982).
respond to the pheromone both to locate a host and potential
mates
(Btrkmshaw.
a possible
than
were
frequently
addrtional
advantage
host
location
Mate selection IS
of females
bemg able to
distmguish between SIgnals from different males.
not those
not always those selected
of therr aggregation
pheromone
host selection
in pairs to females m a matmg
e
1
IS well known
for killing trees (Raffa
by males that have already found a food source.
evolved to attract
of
which IS known to vary in
secure
males
at a distance,
aggregation
males
It has also been observed
others If they are presented
Secondary
until a
land-and-choose
released by closely located males then prehmmary
males (Hodges
males consistently
Such
seems probable that the release of the pheromone
of host
as a preference
secretion,
quantity and quality between
IS found.
IS shown by certam scolytids (Ips spp, Borden,
other
that males and females are
ThIS has been mterpreted
unpublished).
to
density.
which mvolves
behavioural cues causmg
of both males and females
When males releasmg
beetles show a vanable
to particular
source
1993)
released
vanable
determmant
et al.,
attraction
P
male SIgnallers then the Identity
colomsmg males could be an Important
selection
selectd
these
population
land and nutiate test-burrows
in those scolytid beetles responsible
the
mamtenance
or mcreased
behaviour , WIthout response to host volatiles,
ClaSSIC
in male
that have produced
If respondmg
1994)
to different
preferentially
case,
mclude
the beetle to onentate,
SUItable food
differences
Sexually selected traits are notonously
(Andersson,
arena
in this
m determmmg
selected
owes Its ongms
ornaments
Sex-specific
tall and the large antlers of male deer.
process SImilar to those
response
that
tied to
success of individuals
sexually
aggregation
1998)
mdications
acoustic,
are often mstrumental
Perhaps
all
truncatus
biology of P
the reproductive
chemical,
are
pheromone signalhng IS mdeed mtnnately
beetle
would appear to proceed as
field IS fernale-biased,
aggregation
pheromone
histerid
follows. It IS mitiated when beetles take flight m response to
aggregation
that the sex ratio of beetles attracted
nsk as the aggregation
a kairomone
pheromone
The Role and Evolution of Sex and
Aggregation Pheromones in the Life
Histories of Storage Pests
on
Thus the
SIgnal appears not to the behaviour
of the cockroach Nauphoeta ctnerea (Moore et al , 1997),
where vanation m the pheromone signal IS the baSIS for mate
selection on contact
mcrease
Male P. iruncaius
their reproductive
success by manipulation
dIstrIbution through pheromone
make contact,
beetle
signallmg but, once beetles
of tunnellIng
plant hosts
sandWIched
shown
males
form
P
between
truncatus,
paIr
OVIposItmg female and mate repeatedly
(BIrkmshaw,
1998)
bonds
have
storage
WIth an
With that female
a behaVIour they share WIth some speCIes of
down of the aggregation
pheromone
guardmg
to the
and shut
sIgnal, m the presence
that
rely
strategy
reproductive
adult
and a few storage
pheromones
of only the oppoSIte sex
Coleoptera,
the
and non-feedmg
that result m
In contrast,
long-
that feed, such as most of the
on
(aggregatIOn)
of both sexes
IS that the pnonty
locate the oppoSIte sex and mate rather
tunnel system,
Tunnel
noted
30 days)
such as the moths
whIch lead to assemblages
short-lIved
When not copulatmg WIth the female,
1983).
«
lIved adult pests (>30days)
males tend to SIt m the entrance
(KIrkendall,
1990)
IS mediated by (sex)
the attraction
P. truncatus
scolytId
msects,
Coleoptera,
m artifICial
two glass plates,
temporary
(1982,
biology of short-hved
storage
males are selected on other charactenstIcs
Observations
that
Burkholder
would appear to
pheromones
The lOgIC of the
of the adults IS to
than to fmd food.
The female then spreads her eggs m an area perceIved to be
SUItable for the development
of larvae
1980) or after a certam time,
m the absence of host cues,
Just scatters
1791
the
eggs
The
female
(Barrer
and Jay,
IS responsible
for
Proceeduiq« of the 7th Internatwnal
releasing
the sex pheromone.
longer-hved
In contrast,
Working Conference on Stored-proiuct Protectwn - Volume 2
on
the adults of
developmental
stages and produce pheromone
on such hosts. Generally,
that leads to the aggregation.
Most storage
P
the second. However,
Levinson
and
pests fall into
truncatus clearly into
male-produced
aggregation
that
provides
OVIpoSItIon SItes,
benefits
from
aggregation
as
well
The
truncatus,
use
a relatively
the
(Horn)
(Col
Angewandte Schaedmgskunde
65, 8,153 - 157
1982 Aggregation pheromones
In: Bark
eds MItton J. Band
Sturgeon K, B, pp 74 - 139, University
wluch insect
Austm , USA
have stronger
evolved
assocrations
truncatus,
It
would
WIth stored
appear
pheromone m order to attract
the pheromone
between
quality
to locate
different
mates
pheromone
males'
and/or
in those species
that
Males
male
G A.,
to obtam
also respond
IS termmated
truncatus
to gam food resources and mates and aggregatIon
as an unmtentIonal
attractant
beetles
by-product
(ScolytIdae)
courtshIp
where
probably
pheromones
evolved
and male-male
m the
competItIon
beetles.
lead
to
of
mCldentally attracts
Umverslty
In summary,
we suggest
fIrst evolved as sex-attractant
pheromones
the VIews of Levmson and Levmson,
1991
pheromones
W. C,
Press,
Barrer P. M.,
host-food
Sexual
SoClet, 17
U. S. Department
of
Bulletm No
96, 48
of major
R. J
and Pickett,
J A ,
component
of the
male
the role of the
Dawkms R.,
truncatus
(Horn)
(Coleoptera:
Journal of ChemIcal Ecology,
1976. The SelfIsh Gene
17, 4, 789 -
Oxford Umverslty
Press.
Dendy J. , Doble P. , Saidl J A. , SmIth J , and Urono B. ,
selectIOn.
1989 Trappmg the Larger Gram Borer, Prostephanus
truncatus m maIZe fIelds usmg synthetIc pheromones.
Entomologla expenmentahs et apphcata, 50,241- 244
Pnnceton
New Jersey.
1983 A fIeld demonstratIon
fmdmg
gram msects,
1994.
Prmceton,
and
warehouse
Papers on msects affecting stored
DIVISIOnof Entomology,
Prostephanus
1995), and that the
References
Umverslty
and
The Theory of Sex AllocatIon Pnnceton
Hall, D. R. Hodges,
803
Andersson
biology
storage
produced aggregatIon pheromone of the Larger Gram Borer
(m contrast with
of other storage pests in thIS hght.
Reproductive
gram
Pnnceton.
IdentifIcatIon
It would be worth attemptmg
pheromones
m
-52
a mate and
Bostnchldae),
aggregatIon
habitats
et applicata,
The larger gram borer
response by other males IS part of the struggle for surVIval
to remterpret
ecological
expenmentahs
1-10
F. H ,1911
Agriculture,
Cork A.,
that aggregatIon
Press,
products.
(Raffa et
other males
among
Charnov E ,1982
al. , 1993). P. trunca tus would appear to fIt the pa ttern of
a male calhng from ItS food source to attract
1998. Trees
Prostephanus
Journal of the GeorgIa EntomologIcal
Chittenden
and were elaborated
different
1982
E,
(II supplement),
context
further to the benefIt of both senders and receIvers
W.
communication
to a sex
that
sexual
of rrugratmg
Entomologia
Volume 9:
pp. 257 - 285
87,285 - 294.
ThIS IS also conSIdered to be the case m bark
aggregatIon
ongm
collected
Burkholder
IS achIeved
of the response
The
southern Benm.
probable that other males are exploiting the smgal
and pharmacology.
New York
C. , Tchabi A , and Scholz D.,
or m stores?
It IS
once a female has arnved
Pergamon,
Borgemeister
high
to the
does not emit ItS SIgnal for the benefit of other males smce
therefore
biochemistry
Behaviour
as a means of locating a host but the SIgnaller
signallmg
and L I Gilbert
physiology,
discnmmate
of Texas Press,
1985. Agregation pheromones
In: Kerkut,
1985 Comprehensive insect
Borden J H.,
releases
Females respond to
and may
SIgnals in order
hosts
In P.
products
the
females.
a host
which
thesis,
Umweltschutz,
beetles in North Amencan Conifers.
poorly adapted
PhD
G A , and Schulz F A.,
Bostnchidae)
Borden J. H.,
male
pheromone.
storage pest, might well grve some clues as to the means by
aggregation
of Stored
1992. The
mqrescens LeWIS (Col ,
to the pheromone of Prostephanus truncatus
Pflanzenschutz
evoloved
of
Journal
Teretriosoma
of
Histendae)
physical
aggregation
Labonus
response
an
feeding,
as
that sex phermones
pheromones
m P
pheromone
of optunal
SUItable mates
protection ", and presumed
of creating
(Walker)
to OVIPOSIt in
of Leicester , U. K
Boeye J.,
VIewed aggregation
msects as a means'
and
to a sourece of gram odour
Umversity
(1995)
Levinson
Ephestna cautella
of
Phycrtidae ) locate,
16,1,1-7
Birkmshaw L. A , 1998. Mate choice m Prostephanus
truncatus (Horn) (Coleoptera: Bostnchidae): The role of
there are several exceptions hsted by
in storage
ability
Products Research,
Levinson and Levinson (1995)
pheromones
the
response
when feeding
the male produces the pheromone
one or other of these categories:
to
(Lepidoptera:
species feed as adults on the same host as the
behavIOur m several
Journal of Stored Products
of odour-based,
Dendy J , Doble P , Saidl J. A. , SmIth J , and Urono B ,
1991 Tnals to assess the effectIveness
speCIes of stored
Research,
9,3,
pheromone
105-110.
Barrer P M. and Jay E G , 1980. Laboratory
observatIons
1792
mIXtures
for
trappmg
of new synthetIc
Prostephanus
truncat'us (Horn) Products (Coleoptera:
BostrIchidae)
maize stroes.
27,1,69 -74.
Journal of Stored Research,
III
Proceedmas of the 7th Internaiumal
Detmers
H B , 1990
Bedeutung
des
Untersuchungen
holzes
fur
den
MItteilungen
aus
Fortwirchaft.
der
Umversity of Readmg, UK
Markham R H., Wnght V F , and RiOS Ibarra,
zur biologischen
Grossen
Proetephomus truncatus
Biologischen
Worhng Conference on Stored-product Protection - Volume 2
Kombohre
(Bostrvcludae
Bundesanstalt
for
1991
)
truncaius
Land-und
Flight untiation by Proetephan us
truncatus in relation to time of day, temperature, relative
hurrudrty and starvation, Entomologia Expenmentahs
et
Morre P
communication
truncatus
Flight
of the Larger
(Horn)
behavior
(Coleoptera:
TheSIS, Umversity
Bostnchidae
1.,
PhD
mediated by aggregation
truncaius
pheromone
Nzeve
(Horn)
1992.
synthetic
pheromone
R. J,
Prostephanus
Hodges
Bostnchidae
biology
iruncaius
) -
mcreasmg
22,1-14
The
range
Journal
of Stored
pest
with
Products
(Horn)
1994.
Recent
advances
Canberra,
Australia,
17 - 22 Apn11994.
their
the
responses
(Coleoptera:
of
Bostnchldae)
( Coleoptera:
Likhayo P W. ,
Prostephanus
trU1'watus
and SitophLlus zeamaLS
lures m Curculiomdae)
to pheromone
StudIes
Honduras,
Bulletm of En tomologlcal Research,
studIes
on
truncatus
behaViOral
(Horn)
conspeClflcs,
(Coleoptera:
synthetic
TeretrlOsoma
Hlstendae).
mteractiOns
pheromone
mgrescens
of
Natural
Resources
Prostephanus
566 - 569
and
the
(Lewes)
StrategIes
beetles.
(Coleoptera:
Levmson,
structure
A
(Coleoptera:
Press,
Scolytidae
and Levmson,
H , 1995.
of pheromone
ReflectiOns
glands
Phillips
on
-118
Borer,
Dept
Prostephanus
truncatus
Conference
T. W.,
on
J. ,
of the 6th
Stored
Products
17 - 23 April,
1994 1,
and Salom S. M , 1993
of host colomsation
by bark
T. D , and Filip G M. , eds.,
m Comfer Forests.
AcademIC
103 -128
of Applied Entomology,
R
and Herrera
118,
354-
F. L.,
1990
Hlstendae)
(Col
and
: Bostnchldae).
ItS prey
Prostephanus
TropIcal
ScIence,
30,
153 -165
Li L. , 1988. Behavioral ecology and lIfe hIstory evolution m
Grain
pests,
ObservatiOns on the ecology of Teretrwsorna mzgrescens
m storage
LeWIS (Col
PhD ThesIs,
product
In: HIghley E , Wnght E.
Australia,
360
Rees D. P , Rodnguez
truncatus
Larger
stored
1994.
M , Alba-AVIla de A , Ramlrez-Zurbia
Journal
msect speCIes Anzerger for SchodlIngkunde Planzenschutz
(Horn)
of
Interactions
London. pp
Umweltschutz,
68,5,99
16,10,64-70.
(Horn) and S'Ltoph'Lluszearnms
In: Schowalter,
R , 1994.
of the
for the Biological and Integrated
and mechamsms
Ramirez-Martmez,
and
ZoologIcal Journal of the Lmnean SocIety,
and function
Workmg
Beetle-Pathogen
KIrkendall L. R. , 1983. The evolution of matmg systems m
bark and ambrOSIa beetles
traps and
Bulletin
L , WhIte R D. , and Hall DR,
Canberra,
Raffa K. F.,
wIth
predator
Journal of Stored Products Research
Piatypodidae)
77,293 - 352
Institute,
of pheromone-baited
responses
InternatiOnal
m
pp 56.
Gram Borer
to food volatiles.
Protection.
129
truncatus
Banks H J and Champ B. R , eds Proceedmgs
Laboratory
Bostnchidae)
J
of
Motsch,
and
88,2, 131 - 140
and Dobson C C , m press.
Agronomo,
Vlll + P
crevice traps and flight mill
Typentten
Organisation
de
de maiz en el Valle de el
TeSIS de Ingernero
Prostephanus truncatus
synthetiC malze volatiles as lures m crevlce or fllght traps.
Hodges R J.,
truncaius
Prostepluuiue
1991
uses for the Larger
PIke V , Smith
natural
Bostncludae ) m Kenya
Control of NOXiOUSAmmals and Plants,
1998
The
Prostephanus
the
Bostnchidae ) en el ambiente
traditional
report,
International
Volume 2, 929-
Hodges R. J. , Hall D. R. , Mbugua J Nand
179
1,39 - 47
PIke V , 1993b Development
Protection,
934.
the
P,
in MeXICO
Inpubhshed
m the biology and
on Stored Products
pp
1993.
for
(Coleoptera:
relation to pheromone-baited
Research,
on Larger
PhD Tbesis,
PIke. V 1993a
PIke V. , 1993a The behavior of Prostephanus
an
control of Prosteptumus truncatus . In: Proceedmg of the
6th Working Conference
W.,
reservorr
(Coleoptera:
performance
Hodges R J.,
a
Escuela Agncola Panamencana,
of
III Kenya.
College),
J
and Obiero
as
almacenanuento
(Coleoptera:
storage
N.
Zamorano Honduras C A
control
on cockroaches
Hill M G , Chandi E A , cluro C. T. ,
Ramerx,
(Horn)
in response to
and
(Horn)
A destructive
D
Novillo
Tropical SCIence, 32,163 -170
1986
status
Ecological studies
Afncan Crop Science Journal,
Flight behaviro of the
larger gram borer Prostephamu: truncatus
K. F Haynes, and A J
conveys
of London (Impenal
FLO,
truncaius
IS
but not food volatile.
Journal of Stored Products Researcht m press)
Farrell G , and Key G E.,
Odour
environment
and Hodges R. J , 1998.
of Prostephamus
Upwind flight
)
Umversity
and
Ceiba , 32,1 - 90.
N L Reagan-Wallm,
1997.
Nang'ayo,
of Oxford, Green College, pp 227.
Fadarruro H Y , Gudrups
: Bostrichidae ) With an annotated
Gram Borer m Savannah woodlands
Prosteplumus
R.1\1 ,
on Prostephanus
of research
Nature 389,25.
Nang'ayo , F. L. 0 , 1996
and pheromone
Gam Borer,
J ,
Moore,
Apphcata , 75,273 - 277
H. Y , 1995b
(Col
review
updated bibliography
Heft 260,93 p
Fadanuro H Y. , 1995a
Fadanuro
A selective
Scholz D.
1997
Prostephanus
of Pure and ApplIed Zoology,
1793
DIspersal
truncat/iS
and host fmdmg
(Horn)
behaVIor of
(Coleoptera:
Proceeduup: of the 7th International
Bostricludae ).
Germany,
PhD
Thesis,
Umversrty
Borgemeister
pheromone.
53 - 6I.
C.,
truncatus
Infestation
irutiated
by
H -M.,
behavior
of Prostephanus
and Lawson A.,
Bostncludae
):
attack? Journal of Entomology,
et Apphcata , 83,
Vazquez-Ansta
attraction
or
121,261 - 269.
Borer
Research,
Bostnchidae )
32,2,171
Wnght
of attraction
H.,
Olalde-Portugal
Celluloytic
bacteria
Journal
1982.
truncatus
host
selection
of Prostephanus truncatus
to
synthetic
Gram Borer Project,
and
development
1794
kairomones
colorustation
and Stabrawa
Togo
socio-econonuc
of
Resources Institute
pheromone.
and
90 (5),
Annual Review of Entomology,
Akou-Edi D.,
MAP.,
ological
M. E , and Vazquez-A.
digestive
(Coleoptera:
of Economic Entomology
The role of pheromones,
m the
V ,
and Blanco-
446
Larger
Products
R
Infestation
cassava and maize by Prostephanus
Flores-S.
M. , 1993. Flight penodicity
longevity
(Horn)
of Stored
- 18I.
Tigar B J. , Key G. E.,
and
iruucatus
Journal
Smith
behavior of bark beetles
27,411-
pheromone by male Larger Gram
Prostephanus
beetles,
(Coleoptera:
30, 1 , 1 - 8
Prostephanus
of
of maize
of Stored
m the
and allomones
of the effect of females and their residues on
production of aggregation
Journal
Labra A , 1997.
1371-1376.
Wood D L.,
Smith J. L , Cork A. , Hall D. R , and Hodges R. J , 1996
infestation
m Mexico.
R.,
Bostncludae),
(Col.,
random
M.,
pests
97
and Vasquez-A.
Hmojosa R E , Hemandez-Delgadillo
system
Host-fmdmg
(Horn)
products
Products Research,
male-produced
1997b.
truncatus
pnmary
by stored
of maize by
C. , Markham R. H. ,
Scholz D. , Tchabi, A. , Borgemeister
Poehlmg
M. , 1994. Filed and post maturation
Meikle W. G. , Markham R
1997a
Entomologia Expenmentahs
Investigation
et apphcata , 66,91-
Tigar B. J , Key G E , Flores-S. M. E.,
H. , and Poehlmg H. -M.,
Prostephanus
Entomologia expenmentahs
of Hannover,
pp. 146.
Scholz D.,
Applied
Worktng Conference on Stored-product Protection - Volume 2
loss
assessments
reduction
Report R1941
truncatus
strategies.
Typewritten,
A. 1993
of dned
entom
i
for
the
Natural
141p