Thursday 25 June 2015

Aggression Promoting pheromone:Pheromone Resources

21.Aggression Promoting pheromone:

11-cis-vaccenyl acetate (cVA), a male-specific volatile pheromone, robustly promotes male-male aggression in the vinegar fly Drosophila melanogaster (Liming Wang and  David J. Anderson, 2010).


Reference:


Liming Wang and  David, J. Anderson.2010.Identification of an aggression-promoting pheromone and its receptor neurons in Drosophila. Nature.,463:227-231.

Cryptic alarm pheromone:Pheromone Resources

Cryptic alarm pheromone:

Rapidly moving solitary insects can escape the predatory ants whose attack behavior is severely disrupted by the Cryptic pheromone 2-heptanone (Blum et al.,1963).

Reference:

Blum, M.S., Warter, S.L., Monroe, R.S., Chidester, J.C.1963.Chemical releasers of social behaviour. I. Methyl-n-amyl ketone in Iridomyrmex pruinosus. J. Insect Physiol. 9, 881–885.

Alarm pheromone:Pheromone Resources

Alarm pheromone:
Alarm pheromones serve to rapidly disperse a group of insects usually as a response to predation or threat. alarm pheromone, it is generally considered necessary to demonstrate that (i) the chemical(s) is released exclusively under exposure to hazard (e.g. predator attack), (ii) the signal is perceived by conspecifics, and (iii) it induces in the receiving individuals behavioral reactions similar to that induced when directly exposed to the same danger (Wyatt, 2003). Generally, adaptive responses to the reception of an alarm pheromone may be classified as evasive (e.g., receivers flee from the pheromone releaser) or aggressive (receivers move toward the signal and attack or harass the predator). Observed reactions can vary according to the concentration of pheromone released and also with prior experience of the receiver (Howse, 1998).

The alarm pheromones of many arthropods are also used as defensive allomones, activity inhibitors, cryptic alarm pheromones, aggregative attractants, robbing agents, digging agents, trail pheromones, and antimicrobial agents (Blum,1996). 

Alarm pheromones have been identified chiefly in three subfamilies of ants and their distribution appears to be chemosystematically significant. Myrmicine genera emphasize 3-alkanones as alarm releasers, whereas methyl ketones, primarily of terpenoidal origin, are widely utilized as alarm pheromones in the subfamily Dolichoderinae. Formicine species may employ formic acidas an alarm pheromone in addition to the compounds produced in the mandibular and Dufour's glands. The mandibular gland pheromones are chiefly acyclic monoterpene aldehydes (e.g., citronellal) which are relatively low boiling compounds. Higher boiling n-alkanes are produced in the Dufour's glands and may serve as more persistent releasers of alarm behavior. Alarm pheromones as well as the caste-specific pheromones of male bees and ants, probably also serve as defensive products (Blum and  Brand,1972). These kinds of pheromones are usually of short duration and the dispersed individuals usually reform aggregations.  Some individuals release this pheromone to exhibit aggressive behaviour in the presence of predators.  Alarm pheromones have been recorded in the cockroaches, treehoppers, aphids, bedbugs, termites, and social Hymenoptera ( Kerkut and Gilbert, 1985; Blum, 1996). The alram pheromone has also been recorded in fishes (Scholz et al., 2000); amphibians (Scholz et al.,2000) and in mammals like cows and feathered minnos (Mathis et al.,1995).


In Homopterans, so far only in two families the alarm pheromones have been studied.  In membracids the body wall tissue is the source of pheromone (Nault  et al., 1974), whereas, in aphids the cornicles are the source of pheromone (Bowers et al., 1972). In many other aphids alarm pheromones are reported (Wientjens et al.,1873;Bowers et al.,1972;Edwards et al.,1973; Phelan et al.,1976; Pickett and   Griffiths,1980; Gibson and  Pickett,1983).  In Hemiptera, four families are identified possessing alarm pheromone (Blum, 1985).

In Pyrrhocoridae, the 3rd  dorsal abdominal parts; in Cimicidae the dorsal abdominal gland; in Coreidae the metathoracic glands in adults and dorsal abdominal gland of nymphs (Blatt et al., 1998; Leal et al.,1994) have been reported to produce alarm pheromones.
In Pentatomidae,  it is located in adults metathoracic gland (Ishiwatari,1974). In Acanthosomidae,  the alarm pheromone has been reported from dorsal abdominal gland in the  larvae and metathoracic glands in the adults (Maschwitz and Gutmann, 1979).  In Miridae, the metathoracic scent gland is the source of alarm pheromone (Groot et al .,2001). In the Thysanoptera (thrips), the alarm pheromone is recorded from anal fluid produced by the nymphal instars (Teeling et al., 1993).  In Isoptera (termite) the frontal gland of soldiers and homogenial femuli in workers are found to be  the sources of alarm pheromones (Moore, 1968; Farhat and Iqbal, 1980).

The sources of alarm pheromone in ants have been located to eleven; the mandibular glands (Wilson, 1958; Maschwitz, 1964), Pygidal gland (Wilson and Pavan, 1959), Dufour’s gland and poison gland (Maschwitz, 1974) of metaplural gland of workers (Masschwitz, 1964), crushed heads of workers (Duffield et al., 1980). Higher concentration from the heads of aged queens (Francke et al., 1980), small quantities in the abdomens of aged queen (Hughes et al., 2001), whole ants with crushed heads, anal gland  or some other source of pheromones (Hefetz and Lloyd, 1983).

In the family Blattidae of the order Dictioptera, the whole body of the both sexes and glands associated with the 2nd abdominal spiracle are reported to be the sources of alarm pheromone (Brossut, 1983; Rollo et al., 1995). Aggressive behaviour in some of the families of Hymenoptera has also been  demonstrated.  In Vespidae the poison gland of workers and females are located to be the source of alarm pheromone (Maschwitz, 1964).  Similarly, venom reservoir (Manzoli-Palma et al., 1998), pupal odour (Landolt et al., 1999) and heads of workers (Maschwitz, 1984) are identified as the sources of alarm pheromones (Ghent and Gary, 1962; Maschewitz, 1964; Shearer and Boch, 1965).  Even the venom has been identified as the source of alarm pheromone in honeybees (Veith et al., 1978).  Koschewnikow gland is identified as the sources of alarm pheromones (Lensky et al., 1991).  Cassier et al. (1994) reported sting sheets as a source of alarm pheromone in honeybee. Most Alarm pheromones likely have evaloved from compound originally having other functions. Specially it has been proposed that Alarm pheromone may evalve from  chemicals used for defense against predators or from compounds released upon injury (Wyatt, 2003).

The aggregation pheromone of  Wisconsin Ips pini contains ipsdienol, which is (+)  and (-) enantiomers, and lanierone, and the pheromone  of I. grandicollis contains ipsenol. By contrast,  -pinene consistently enhanced attraction by all three  predators to the pheromones of their Ips prey (Nadir Erbilgin  and Kenneth F Raffa, 2001).For a detailed review of the alarm pheromone, the research work of Blum (1985) may be consulted. 

Similarly 2-heptanone and 2-heptanol are known as ‘alarm’ or repellent substances in many hymenopterans, including honey bees (Free, 1987; Wongsiri et al., 2006) and several ant species (Vander Meer et al., 1998). A recently accepted patent (US patent number 6,071,973) by Vander Meer et al.(2000). In  many ants and termites, the glandular sources of trail and alarm pheromones have been reported (Kaib, 1999; Kaib, 2000;Wyatt, 2003).

References:

Blatt, S. E., Borden, J. H., Pierce, H.D. Jr., Gries, R. and  Gries, G. 1998. Alarm pheromone system of the western conifer seed bug, Leptoglossus occidentalis. Journal of chemical ecology (USA).,24(6):1013-1031.
Blum, M. S. 1996. Semiochemical parsimony in the Arthropoda. Annual review of entomology, 41(1):353-374.
Blum, M.S. 1985.  Alarm Pheromones in Comprehensive Insect Physiology Biochemistry and Pharmacology (eds. G.A. Kerkut and L.I.Gilbert) Pregman Press,Oxford,9:193-224.
Blum, M.S. 1985.  Alarm Pheromones in Comprehensive Insect Physiology Biochemistry and Pharmacology (eds. G.A. Kerkut and L.I.Gilbert) Pregman Press,Oxford,9:193-224.
Bowers, W. S., Nault, L. R., Webb, R. E.and   Dutky, S. R.1972.Aphid alarm pheromone. Isolation, identification, synthesis. Science, 177:1121-1122.
Cassier, P. Tel-Zur, D. and Lensky, Y. 1994. The sting sheaths of honey bee workers (Apis mellifera L.): structure and alarm pheromone secretion.Journal of Insect Physiology.,40(1):23-32Cole, LK. 1974. Antifungal, Insecticidal, and Potential Chemotherapeutic Properties of Ant Venom Alkaloids and Ant Alarm Pheromones.PhD Thesis. University of Georgia, Athens.
Edwards, L. J., Uden, P. and   Kislow, C. J.1973. Trans-b-farnesene, alarm pheromone of the green peach aphid, Myzus persicae.Nature.,241:126−127.
Farhat, A. and Iqbal, Q.J.1980. Extraction and biological evaluation of the trail pheromone of the termite, Odontotermes obesus (Rambur). Biologia, Pakistan.,26(1-2):183-185.
Free, J. B. 1987.Pheromones of Social Bees. New York: Cornell University Press.
Ghent Ralph L. and Gary Norman E.. 1962.A Chemical Alarm Releaser in Honey Bee Stings (Apis mellifera L.).Psyche 69:1-6.

Gibson, R. W. and Pickett, J. A. 1983.Wild potato repels aphids by release of aphid alarm pheromone. Nature., 302:608-609.

Groot, A.T., Drijfhout, F.P., Heijboer, A., Beek,T.A van and Visser, J.H. 2001. Disruption of sexual communication in the mirid bug Lygocoris pabulinus by hexyl butanoate. Agricultural and Forest Entomology.,3(1):49-55.

Ishiwatari, T. 1974. Studies on the scent of stink bugs (Hemiptem Pentatomidae) I. Alarm pheromone activity. Appl. Entomol. Zool. 9: 153-158.

Kaib, M. 1999.Termites. In Pheromones of Non-Lepidopteran Insects Associated with Agricultural Plants (ed. J. Hardie and A. K. Minks), pp. 329-353. Oxon: CAB International.

Kaib, M. 2000.Chemical signals and communication in termites: a review. Mitt. Dtsch. Ges. Allg. Angew. Entomol. 11:211-218.

Kerkut, G. A., and Gilbert, L. I. (eds.).1985.Comprehensive Insect Physiology, Biochemistry and Pharmacology,Pergamon Press, Elmsford, NY.
Landolt, P.J., Reed, H.C. and Heath, R.R. 1999. An alarm pheromone from heads of worker Vespula squamosa (Hymenoptera: Vespidae).Florida Entomologist., 82(2):356-359.
Leal, W.S., Panizzi, A.R. and Niva, C.C. 1994. Alarm pheromone system of leaf-footed bug Leptoglossus zonatus (Heteroptera:Coreidae). Journal of Chemical Ecolog.,1994, 20(5):1209-1216.
Lensky, Y., Cassier, P., Rosa, S. and Grandperrin, D. 1991. Induction of balling in worker honeybees (Apis mellifera L.) by "stress" pheromone from Koschewnikow glands of queen bees behavioural, structural and chemical study.Comparative-biochemistry-and-physiology,Comparative physiology(USA). v., 100(3):585-594.
Manzoli-Palma,M.F.,Gobbi,N, Palma,M.S.1998.Alam pheromone and influence of  pupal odour  on  the aggressiveness od  Palybia paulista (Ihering) (Hymenoptera :Vespidae), Journal of Venomus Animal and Toxins,4 :61-69.
Maschwitz, U and Gutmann, G.1979. Trail and alarm pheromones in the speckled brooding bug Elasmucha grisea (Heteroptera: Acanthosomidae). Insectes Sociaux.26(2):101-11.
Maschwitz, U. 1964.  Gefahrenalarmstuffe und Gefahrenalarmierung bei sozialen Hymenoptera. Z. Vergl. Physiol. 47 : 596-655.
Maschwitz, U. 1974. Vergleichenda Untersuchunger zur Funktion der Ameisenmetathorakaldruse.  Oecologia, 16 : 303-310. 
Maschwitz, U.1984. Alarm pheromone in the long-cheeked wasp Dolichovespula saxonica (Hym. Vespidae).Deutsche Entomologische Zeitschrift.31(1-3): 33-34.
Moore, B.P. 1968.  Studies on the chemical composition and function of the cephalic gland secretion in Australian termites. J. Insect Physiol. 14 : 33-39.
Nault, L.R. , Wood, T.K. and Goff, A.M.1974.  Treehoppers (Membracidae) alarm pheromones.  Nature,  249:387- 388.
Phelan, P. L., Montgomery, M. E., and  Nault, L. R. 1976. Orientation and locomotion of apterous aphids dislodged from their hosts by alarm pheromone. Annals of the Entomological Society of America,69(6):1153-1156.
Pickett, J. A. and   Griffiths, D. C.1980.Composition of aphid alarm pheromones.J. chem. Ecol.,6:349−360.
Scholz, N.L., Truelove, N.K., French, B.L., Berejikian, B.A., Quinn, T.P., Casillas, E., Collier,T.K.2000. Diazinon disrupts antipredator and homing behaviors in chinook salmon (Oncorhynchus tshawytscha). Canadian Journal of Fisheries and Aquatic Sciences.57(9):1911-1918.
Shearer, D.A., and Boch, R. 1965. 2-heptanone in the mandibular gland secretion of the honey-bee. Nature. 204:530.
Teerling, C.R., Pierce, H.D. Jr., Borden, J.H. and Gillespie, D.R. 1993. Identification and bioactivity of alarm pheromone in the western flower thrips, Frankliniella occidentalis. J Chem Ecol.,19(4):681-697.
Vander Meer R.K. and Alonso L.E., 1998.Pheromone directed behavior in ants.in : Pheromone communication in social insects : ants, wasps, bees, and termites Vander Meer R.K., Breed M.D. and Winston M.L. K.E. Espalia (eds.), Westview Press : 159-192.
Veith, H.J., Weiss, J., Koeniger, N.1978. A new alarm pheromone (2-decen-1-yl-acetate) isolated from the stings of Apis dorsata and Apis florea (Hymenoptera: Apidae).  Experientia.34(4): 423.
Wientjens,W.H.J.M.,Lakwijk,A.C.and Van der Marel, T.1873.Alarm pheromone of grain aphids. Experientia.,29:658−660.
Wilson, E.O. and Pavan, M. 1959.  Glandular sources and specificity of some chemical releases of social behavior in delichoderine ants. Psyche 66 : 70-76.
Wilson, E.O.1958. A chemical releaser of alarm and digging behavior in the ant, Pogonomyrmex badius (Latreille).  Psyche 65 : 41-51.
Wongsiri, S., Suwannapong, G., Srisook, N. and Hepburn, R. 2006. Pheromones of Asian Honeybees (Apis andreneformis, Apis cerana, Apis dorsata and Apis  florea). In XV Congress IUSSI Proceedings, 30 July-4 August, 2006. Washington DC: IUSSI.

Wyatt, T. 2003.Pheromones and Animal Behaviour. Cambridge: Cambridge University Press.

Wednesday 24 June 2015

Aggression inhibiting pheromones:Pheromone Resources

Aggression inhibiting pheromones:

In mice, Jones and  Nowell (1975)reported  that the coagulating glands are also shown to be the source of an aggression-inhibiting pheromone. The common source of these two factors suggests that they may be the same pheromone exerting different effects under different conditions. The territorial implications of both the aversive pheromone and the aggression-inhibiting pheromone are discussed (Jones and  Nowell, 1973).


Ovary inhibiting pheromone:
Queen Retinue Pheromone also acts as a primer pheromone by physiologically inhibiting the ovary development of worker bees. An essential component of QRP, 9-oxo-(E)-2-decenoic acid, acts as a long-distance sex pheromone (Trhlin and Rajchard,2011).

Puberty inhibiting pheromone:
The onset of puberty in female house mice (Mus musculus domesticus) can vary from the age of 4 wk to 8 wk as a function of male acceleratory or female inhibitory urinary pheromones. In field studies, puberty inhibition has been demonstrated directly with natural and experimental increases in population density(Rjohn G Vandenbergh, 1987).

Courtship-inhibiting pheromone:
Gregarious mature males of the desert locust (Schistocerca gregaria) emit a courtship-inhibiting pheromone continuously to repel rivals. This signal evokes a strong response from males with recent experience of mature females. However, if males have been female deprived for some time, they start to ignore the pheromone and attempt to usurp females that are guarded by males. The probability and intensity with which males struggle for an occupied mate was found to depend on the time previously spent without a female (Karsten Seidelmann, 2006).
Swarming Inhibiting Pheromone:

Footprint pheromone -The footprint substances are excreted by workers, drones and queens.

1.Worker foot-print substances areused fororientation to hive entrance, attraction to food source, survival of isolated workers, acceptance of grafted larvae in queen cups, drone mating flight (Butler et al.,1969).

(R,E)-(−)-9-Hydroxy-2-enoic acid (9-HDA) promotes stability of a swarm, or a "calming" influence Footprint pheromone. This pheromone is left by bees when they walk and is useful in enhancing Nasonov pheromones in searching for nectar. In the queen, it is an oily secretion of the queen's tarsal glands that is deposited on the comb as she walks across it. This inhibits queen cell construction (thereby inhibiting swarming), and its production diminishes as the queen ages.


Juvenile mouse pheromone inhibits sexual behaviour:

A juvenile pheromone produced by young mice before puberty, termed exocrine-gland secreting peptide 22 (ESP22). ESP22 is secreted from the lacrimal gland and released into tears of 2- to 3-week-old mice. Upon detection, ESP22 activates high-affinity sensory neurons in the vomeronasal organ, and downstream limbic neurons in the medial amygdala. Recombinant ESP22, painted on mice, exerts a powerful inhibitory effect on adult male mating behaviour, which is abolished in knockout mice lacking TRPC2, a key signaling component of the vomeronasal organ (David M. Ferrero et al.,2013).

Pheromone inhibition of virulence factor :

The extracellular signal of the quorum-sensing system is a thiolactone-containing peptide pheromone, whose sequence varies among the different staphylococcal strains. We demonstrate that a synthetic Staphylococcus epidermidis pheromone is a competent inhibitor of the Staphylococcus aureus agr system. Derivatives of the pheromone, in which the N-terminus or the cyclic bond structure was changed, were synthesized and their biological activity was determined. The presence of a correct N-terminus and a thiolactone were absolute prerequisites for an agr-activating effect in S. epidermidis, whereas inhibition of the S. aureus agr system was less dependent on the original structure (Michael Otto et al.,1999).

Pheromone Cross-Inhibition:

Cross-inhibition by quorum-sensing pheromones betweenStaphylococcus aureus and Staphylococcus epidermidis was investigated using all known S. aureus agr pheromone subgroups. All S. aureus subgroups were sensitive towards the S. epidermidis pheromone, with the exception of the recently identified subgroup 4. The subgroup 4 pheromone was also the only S. aureus pheromone able to inhibit the S. epidermidis agr response (Michael Otto et al.,2001).

Michael Otto, Hartmut Echner,Wolfgang Voelter and Friedrich Götz.2001. Pheromone Cross-Inhibition betweenStaphylococcus aureus and Staphylococcus epidermidis. Infect. Immun.  69(3):1957-1960.


Michael Otto, Roderich Süßmuth, Cuong Vuong, Günther Jung, Friedrich Götz. 1999.Inhibition of virulence factor expression in Staphylococcus aureus by the Staphylococcus epidermidis agr pheromone and derivatives, FEBS Leetrs, 450(3): 257–262.




Oviposition-inhibiting pheromone:



Ferguson A.W., Ziesmann J., Blight M.M., Williams I.H., Wadhams L.J., Clark S.J., Woodcock C.M. and Mudd A.1999: Perceptiom of oviposition deterring pheromone by cabbage weevil (Ceutorhynchus assimilis). J. Chem. Ecol.,25: 1655-1670.

     Oviposition deterrent Pheromone:

The  braconid wasp (Diachasma alloeum) attacks two species of fruit-parasitic flies in the genus Rhagoletis. Female wasps lay a single egg into a second or third instar fly maggot.Following oviposition, female wasps press and drag their ovipositor across the fruit surface with this putative oviposition-deterring pheromone (Stelinski et al., 2007).
Prokopy, R., 1981b.  Oviposition-deterring pheromone system of apple maggot flies.  In E. R. Mitchell, ed. Management of insect pests with semiochemicals.  New York:  Plenum Press, pp. 477-497.

Stelinski L. L.,, R. Oakleafand C. Rodriguez-Saona. 2007. Oviposition-deterring pheromone deposited on blueberry fruit by the parasitic wasp.Diachasma alloeum Behaviour 144: 429-445.


Antibiotic-induced inhibition of pheromone:

Ingestion of diet containing streptomycin inhibited the conversion of myrcene, a host plant terpene, to the male-specific pheromones ipsenol and ipsdienol in Ips paraconfusus. Synthesis of cis-verbenol, which is not a sex-specific pheromone, from the host plant terpene (-)-alpha-pinene and other metabolites from these two terpenes is,as not inhibited by the antibiotic (Byers and Wood,1981).
adenylate cyclase inhibiting pheromone:

Cells of the a mating type, serves to synchronize the opposite mating type (a cells) at GI as a prelude to fusion of the two cell types. It was found that, in vitro, A factor inhibited the' membrane-bound adenylate cyclase Of these cells in a dose-dependent manner. Moreover, one
Class (steS) of a cell mutants that grow normally at either 230 or 340C but that are unable to respond to a factor or to mate at the higher temperature possessed an adenylate cyclase activity that was not inhibited by a factor at 340C but was fully sensitive to inhibition at 230C (Hans Liao and Jeremy Thorner,1980).

Hans Liao and Jeremy Thorner.1980. Yeast  mating pheromone a factor inhibits adenylate cyclase (cyclic AMP/Saccharomyces cerevisiae/plasma membranes/mutants/peptides).Proc. Natl.Acad.Sci.USA 77(4):1898-1902.





Byers, J.A. and Wood, D.L. 1981. Antibiotic-Induced Inhibition of Pheromone Synthesis in a Bark Beetle. Science 213:763-764.


References:

Butler, C.G. and D.H. Calam. 1969. Pheromones of the honeybee—The secretion of the Nassanoff gland of the worker.Journal of Insect Physiology.,15(2): 237-244.
David M. Ferrero, Lisa M. Moeller, Takuya Osakada, Nao Horio, Qian Li, Dheeraj S. Roy, Annika Cichy, Marc Spehr, Kazushige Touhara and  Stephen D. Liberles.2013. A juvenile mouse pheromone inhibits sexual behaviour through the vomeronasal system, Nature, 502:368–371.
Jones, R.B.  and N.W. Nowell. 1973.The coagulating glands as a source of aversive and aggression-inhibiting pheromone(s) in the male albino mouse. Physiology and Behavior.,11(4):455-462.
Jones, R.B. and  Nowell, N.W.1975. Effects of clean and  soiled sawdust substrates and of different urine types upon aggressive behavior in male mice. Aggressive Behavior.,1:111-21.
Karsten Seidelmann.2006. The courtship-inhibiting pheromone is ignored by female-deprived gregarious desert locust males, Biol Lett. 2(4): 525–527.
Rjohn G. Vandenbergh. 1987. Regulation of puberty and its consequences on population dynamics of mice  integrative and comparative biology . 27(3): 891-898.
Trhlin,M and J.Rajchard.2011. Chemical communication in the  honeybee (Apis mellifera L.): a review.Veterinarni Medicina, 56, 2011 (6): 265-273.




Aggression pheromone as alarm pheromone:Pheromone Resources

Aggression pheromone as alarm pheromone:

Alarm pheromone information is transmitted, via projection neurons, in ants to the lateral horn and the calyces of the mushroom body of the protocerebrum. These neurons may participate in the control of aggressive behavior, which is sensitized by alarm pheromones and is triggered by non-pheromonal sensory stimuli associated with a potential enemy (Mizunami et al.,2010).

Alarm pheromone as Aggression pheromone:

Honey bees (Hymenoptera: Apidae, Apini) produce alarm pheromone in the sting gland and a pheromone that has sometimes been described as an alarm pheromone in their mandibular glands, but which does not galvanize colony aggression (Vallet et al., 1991; Couvillon et al., 2010). In the highly social bees (honey bees and stingless bees), alarm pheromones can increase aggression and flight activity at the nest entrance, a defensive response (Boch and Shearer, 1971; Roubik, 1989). Honeybees became adapted to synthetic alarm pheromone components dispensed within their hives and were less inclined to sting. The reduction in the stinging response of honeybee colonies which were adapted to 6 or 8 alarm pheromone components was no more than in colonies adapted to 3 components only, synthetic alarm pheromones to reduce aggression (Free, 1988).
Alarm pheromones play an important role in social insects and enhance collective fitness by providing information about dangers such as predators (Billen and Morgan, 1998). For example, honey bees can use alarm pheromones to co-ordinate colony defense (Free, 1987; Pirk et al., 2011). The behavioral responses can be classified  into (1) initial preparatory phase of alarm behavior, (2) alarm behavior and (3) aggression against a potential enemy (Makoto Mizunami et al.,2010). Aggressive behaviour increases with age when the honey bees are exposed to alarm pheromone ( Alaux et al.,2009). Not only it it increases the aggressive behaviour,but also increases metabolism in honey bees (Southwick  and Moritz, 1985) and higher cytochrome c oxidase activity in rodents and lizards (Sakata et al.,2005). Intrestingly, a reduced metabolism and aggression has been reported for some parts of the human brain (Anckarsa¨ter, 2006). Melanocortins in the preputial glands can alter the excretion of aggression-modifying pheromonesn (Caldwell  and Lepri, 2002.).

The ‘one gland – two functions hypothesis’ has developed (Kerr and da Cruz, 1961)  in meliponine communication. If one assumes that mandibular gland secretions do indeed induce both scent trail following to distant food sources in newly recruited worker bees and defensive/aggressive behaviour in the same workers near the nest. Almost all workers attracted to target 1 became its aggressors, probably due to the presence of alarm pheromones. Thus, pupal odor seems to act as a chemical signal to indicate the presence of pupae in the nest, attracting the workers which would care for and defend the pupae against predators. Although pupal odor does not elicit alarm behavior, it appears to interact with alarm pheromones by potentiating their effects (Manzoli-Palma et al.,1998).



References:

Alaux, C., Sinha, S., Hasadsri, L., Hunt, G. J., Guzmán-Novoa, E., DeGrandi-Hoffman, G. and Robinson, G. E. 2009.Honey bee aggression supports a link between gene regulation and behavioral evolution. Proceedings of the National Academy of Sciences, 106(36):15400-15405.
Anckarsa¨ter, H.2006. Central nervous changes in social dysfunction: Autism, aggression, and psychopathy. Brain Res Bull 69:259–265.
Billen, J. and Morgan, E. D. 1998.Pheromone communication in social insects: sources and secretions. In Pheromone Communication in Social Insects (ed. R. K. Vander Meer, M. D. Breed, K. E. Espelie and M. L. Winston), pp. 3-33. Boulder, CO:Westview Press.
Boch, R. and Shearer, D. A. 1971. Chemical releasers of alarm behaviour in the honey-bee, Apis mellifera. J. Insect Physiol. 17:2277-2285.
Caldwell, H.K. and Lepri, J.J. 2002.Disruption of the fifth melanocortin receptor alters the urinary excretion of aggression-modifying pheromones in male house mice. Chem. Senses, 27, 91–94.
Couvillon, M. J., Barton, S. N., Cohen, J. A., Fabricius, O. K., Kärcher, M. H.,Cooper, L. S., Silk, M. J., Helanterä, H. and Ratnieks, F. L. W. 2010. Alarm pheromones do not mediate rapid shifts in honey bee guard acceptance threshold. J.Chem. Ecol. 36, 1306-1308.
Free, J. B. 1987. Pheromones  of  Social Bees. London: Chapman and Hall.
Free, J. B. 1988. Adapting honeybees (Apis mellifera L.) to synthetic alarm pheromones to reduce aggression, Journal of Apicultural Research, 27(4):227-229.
Kerr, W. E. and da Cruz, C. C. 1961.Funções diferentes tomadas pela glândula mandibular na evolução das abelhas em geral e em Trigona (Oxytrigona) tataira emespecial. Rev. Bras. Biol. 21, 1-16.
Kiyokawa, Y., Kikusui, T., Takeuchi, Y.and  Mori, Y. 2004. Alarm pheromones with different functions are released from different regions of the body surface of male rats. Chemical senses, 29(1), 35-40.
Makoto Mizunami, Nobuhiro Yamagata and Hiroshi Nishino.2010. Alarm pheromone processing in the ant brain: an evolutionary Perspective,Frontiers in Behavioral Neuroscience,4:1-9.
Manzoli-Palma, M. F., Gobbi, N. and  Palma, M. S. 1998.Alarm pheromones and the influence of pupal odor on the aggressiveness of Polybia paulista (Ihering)(Hymenoptera: Vespidae). Journal of Venomous Animals and Toxins,4(1):61-69.
Mizunami, M., Yamagata, N. and Nishino, H.2010. Alarm pheromone processing in the ant brain: an evolutionary perspective. Front. Behav. Neurosci.4:28.
Roubik, D. W. 1989. Ecology and Natural History of Tropical Bees. New York, NY:Cambridge University Press.
Sakata, J.T., Crews, D., Gonzalez-Lima, F.2005.Behavioral correlates of differences in neural metabolic capacity. Brain Res Brain Res Rev 48:1–15.
Schorkopf, D. L. P., Hrncir, M., Mateus, S., Zucchi, R., Schmidt, V. M. and  Barth, F. G. 2009.Mandibular gland secretions of meliponine worker bees: further evidence for their role in interspecific and intraspecific defence and aggression and against their role in food source signalling. Journal of Experimental Biology, 212(8):1153-1162.
Southwick, E.E. and Moritz, R.F.A.1985.Metabolic response to alarm pheromone in honey bees. J Insect Physiol 31:389–392.

Vallet, A., Cassier, P. and Lensky, Y. 1991. Ontogeny of the fine structure of the mandibular glands of the honeybee (Apis mellifera L.) workers and the pheromonal activity of 2-heptanone. J. Insect Physiol. 37:789-804.

Saturday 20 June 2015

Use of Pheromones in managing insects of Storage:Pheromone Resources

Use of Pheromones in managing insects of  Storage
V Nandagopal1, Anand Prakash2 and T. V. Prasad3
1Ex. Senior Scientist, CRRI, Cuttack,Odhisa
2HOD, Crop Protection, CRRI, Cuttack,Odhisa
3Senior Scientist, NBPGR, Pusa, New Delhi-110012

Two major groups of the most economically important post-harvest insect pests: Coleoptera (beetles) and Lepidoptera (moths and butterflies). Several Coleopteran and Lepidopteran species attack crops both in the field and in store. Crop damage by Lepidoptera is only done by the larvae. Several lepidopteran larvae entangle the feeding media through silky secretion which turns products into entwined lumps. In the case of Coleoptera, both larvae and adults often feed on the crop and the two stages are responsible for the damage.

Post-harvest insect pests may be primary, i.e. able to attack intact grains such as the genus Sitophilus, while others are secondary pests, attacking already damaged grains or grain products such as the genus Tribolium. The following is a list of the most common post-harvest and storage pests, their biology, distribution and common host plants.

Coleoptera
Coleoptera is the largest order of insects and contains the most common and important stored product pests. Adults have their forewings modified as hard elytra. Beetles inhabit a wide variety of habitats and can be found almost everywhere. Those associated with stored products exhibit different behavioural types; some are primary and secondary pests feeding directly on the product, others are general scavengers, fungus feeders, wood borers or predators of other insects. arvae lack the presence of prolegs (abdominal legs) and only possess true legs on the three thoracic segments. Larvae of a few species may also lack true legs, e.g. Sitophilus spp.

Curculionidae (Snout Beetles)
This is a large group of beetles that contains some of the most serious crop and stored grain pests. Members of this family are characterised by the form of the snout (rostrum) which is elongated in most species. This family contains the most destructive stored grains pests in the world.

Rice Weevil: Sitophilus oryzae (L.) (=Calandraoryzae L.)
Maize Weevil: Sitophilus zeamais Motsch. (=Calandra zeamais Motsch.)
Granary Weevil: Sitophilus granarius (L.)

The first two species are major primary pests that have a virtually cosmopolitan distribution throughout the warmer parts of the world. The rice weevil (S. oryzae) mainly attacks rice and wheat in stores, while S. zeamais is a serious primary pest of stored maize. However, both species are able to develop on all cereals, dried cassava and other processed food products. The two species are morphologically identical. In Europe, the two species are replaced by the granary weevil, S. granarius, which is wingless and can be distinguished by the sculpturing on the prothorax and elytera.

Natural history:
The life cycle and damage caused by both S. oryzae and S. zeamais are similar. However, S. zeamais is a little larger (5 mm in length) and a very active flier. Infestation usually starts in the field and later continues in the store. Both species are capable of inhibiting reserved breeding grounds near the threshing floors that are normally full of plant residues, where the population builds up in before moving to granaries. Adult females chew grains creating a small hole in which they lay eggs and then seal the hole with a secretion. The optimum temperature for oviposition is around 25oC and at grain moisture contents of over 10 percentage (Brich, 1944). Larvae tunnel in grains and are responsible for most of the damage. Pupation takes place inside the grain and adults chew their way out through the outer layer of the grain. Adults live for 5-6 months depending on the temperature and humidity of grains (Kuschel, 1961; Giles, 1969; Mound, 1989).   S. oryzae adult females can lay more than 500 eggs during their lifetime. The optimal temperature for development is 300C with maximum oviposition taking place at 18 percentage humidity. The rice weevil can live without food for 6-32 days depending on temperature. This species is highly affected by changes in temperature; all stages die in about a week at 00C. On the other hand, S. zeamais tolerates lower temperatures than S. oryzae and can live for 37 days at 00C (Floyd, & Newsom, 1959; Stoyanova, 1984; Zewar, 1993).

Tribolium confusum J. du Val (Tenebrionidae- Darkling beetles)
This is a large and varied group of insects that contains more than 10,000 species of which about 100 are associated with stored products. Most of the tenebrionids are black or dark brown in colour and mainly phytophagous. Adults are characterised by the tarsi of the hind leg with only four segments. Infestation by these beetles results in an unappealing smell due to the secretion of benzoquinones from abdominal glands. The following tenebrionids are serious secondary pests of stored grains and flour.

Confused flour beetle:
These two species are probably the most common secondary pests of all plant commodities in store throughout the world. Several other species of Tribolium are occasional minor pests and can be found in almost every store containing infested cereals or cereal products, specially in tropical and sub-tropical climates. Both species attack maize, wheat, flour and other foodstuffs, but T. confusum does not seem to be as common as T. castaneum in tropical climates (see Hill, 1987; Mills & White, 1994). Members of genus Tribolium are known to produce toxic quinones which contaminate flour and flour products (Gorham, 1989). Damage is done by both larvae and adults specially to broken or damaged grains.

Natural history:
T. castaneum adult females lay small, cylindrical, white eggs scattered in the product. At an optimum temperature of 32.50C, females lay up to 11 eggs daily. Larvae are yellowish with a pale brown head, and they live inside grains until pupation. Adults are about 3-4 mm long and can live for a year or more. Females are highly fecund and able to lay a maximum of 1000 eggs during a lifetime, with 400C and 220C as upper and lower limits for development. This species is also highly tolerable to humidity as low as 11 percentage. Adults are highly adapted to feed on a very wide range of commodities and perfect colonizers of new habitats. In tropical conditions, this species is dominant to T. confusum (Howe, 1962; Dawson, 1977).
The confused flour beetle, T. confusum, is often confused with T. castaneum but they can be separated using the last three segments of the antenna which are much larger than the rest in T. castaneum and forming a club, while the last five segments in T. confusum gradually enlarge towards the tip. Just like T. castaneum, the confused flour beetle develops in crushed grain products and a constant inhabitant of flour mills specially in the temperate regions of the world. In contrast to T. castaneum, this species is not able to fly, but has a long life span that can reach three years under moderate climatic conditions (25-300C) (see Sokoloff, 1972; 1974; 1977).

Yellow mealworm beetle: Tenebrio molitor L.

Natural history:
Tenebrio beetles are black or dark brown and they feed as larvae and adults on grain products. T. molitor is an important post-harvest pest and occurs spread all over the world. Adults are elongate, 16 mm long, and active fliers. Females can lay up to 600 eggs during its lifetime. Larvae firstly eat the germs of stored grains and can feed on a wide variety of plant products such as ground grains, flour, tobacco and foodstuffs. Larvae are very voracious and highly resistant to low temperature; they can remain alive for 80 days at -50C.

Other tenebrionids are less common polyphagous pests around the globe such as T. destructor, T. madens and Palorus depressus.

Bostrichidae (Branch and twig borers)
Members of this family are elongate with the head bent down ventrally to the thorax. Adults are characterised by rasp-like hooks on the pronotum. Most of the species are borers in wood or roots. Wood boring activities of these beetles may weaken timbers or wooden walls of the stores. This family contains two serious stored grain pests:

The lesser grain borer (R. dominica) attacks a wide range of stored cereals. It can be found attacking cassava, flour and other cereal products and is also able to attack rough rice grains. The pest originated from South America, but is now found in all the warmer parts of the world. This species is a serious pest in Australia, from where it was carried to the USA and other parts of the world during World War I. Adults of this species are tiny dark beetles, 2-3 mm in length, and are very voracious with a long life span. Females may continue to lay eggs for four months and are able to lay up to 500 eggs at 340C. They feed externally on grains and lay eggs on their surface. Larvae feed either externally or inside the grain and pupation takes place within the eaten grain. Larval development is relatively faster when fed on whole grains than on flour. Both adults and larvae eat the endosperm leaving powdered grains. This dust can accumulate on the walls of the warehouses and it is a sign of high infestation. Though are not common on pulses, adults are able to breed in grains that are too dry for fast development of Sitophilus. At 340C, development is possible on grains with moisture contents as low as 9 percentage, and they can daily destroy grains equal to their body weight (see Birch, 1945; Fisher, 1950; Aitken, 1975).


Bruchidae (Seed beetles)
Most bruchids are short, stout-bodied beetles with a short forewing not reaching the tip of the abdomen. Adults are characterised by their compact hairy bodies and relatively long antennae. Larvae of most species feed inside seeds and some develop in stored dry grains or legumes. All bruchids are phytophagous with most species able to avoid feeding on seed covers that contain toxins. This family contains several important field and stored crop pests.

Cowpea weevil: Callosobruchus maculatus (Fabricius)
This is an important pest that mainly attacks beans of various species, and can alternatively attack other pulse crops (Lienard and Seck, 1994). This species originated in Africa but is now found all over the tropics and sub-tropics. Adults are 2-4 mm, brownish with black markings. They have a short life span of about 12 days and do not feed. Two forms of this species have been identified; the active (flying) form and the flightless form. The flying form disperses and colonises cowpea fields. Adult females lay about 100 eggs glued to the seed surface or to pods. Larvae tunnel inside the seed where the entire development takes place. In the store, the normal form continues to reproduce until the end of the storage season. The flying form appears again in response to disperses to new locations. This species causes major problems in Nigeria and Niger, where most of Africa's cowpeas are produced (Alebeek, 1996). Other species such as C. rhodesianus and C. subinnotatus may also be important in some parts of Eastern and Central Africa (Gillon et al., 1992; Giga et al., 1993).

American bean weevil: Acanthoscelides obtectus Say (Bruchus obtectus Say).
This species is widely distributed in Africa, Central and South America, New Zealand, USA and Southern Europe. A. obtectus exhibits high tolerance to varied degrees of temperature, thus, it is found in cool highland areas as well as the warmer parts of the tropics. It mainly attacks beans of various types and other pulse crops. Adults are grey and oblong in shape, with the body covered by yellowish green hairs. Females are almost twice as large as males. Infestation starts in the field when females lay eggs on the mature beans in plant pods. Larvae are tiny with strong mandibles and feed inside the seeds where life cycle is completed. Adults exit the seed through

Cucujidae (Flat bark beetles)
Members of this family are small flattened beetles, mostly found under the bark of trees or in tunnels made by other beetles. This family contains one common pest of stored grains.

Red rust grain beetle: Cryptolestes ferrugineus (Stephens)
Adults of this species are oblong flattened small beetles (1.5-2 mm long), with the head and prothorax relatively big and conspicuous. C. ferrugineus is a widespread secondary pest of stored grains, specially in the humid tropics. The genus Cryptolestes was reported to be of economic importance towards the end of the maize storage season in Togo (Pantenius, 1988). However, it might not be as serious as other pests in stores, often following an infestation by other insects. It usually attacks the germs of broken or cracked grains thus reducing germination. Other species such as C. pusillus (Schonherr) and C. pusilloides (Steel and Howe) are common in humid areas of the tropics (Banks, 1979).

Silvanidae: This family was formerly included in Cucujidae. It includes two important species:
Saw-toothed grain beetle: Oryzaephilus surinamensis (L), recognized by the toothed lateral margins of the pronotum.
Merchant grain beetle: Oryzaephilus mercator (Fauvel), which is found in association with O. surinamensis.

Both species are virtually cosmopolitan and they infest a wide variety of stored grains, processed foodstuff and other food products. They are mainly secondary on stored products following more destructive primary pests. However, O. surinamensis prefers cereal products while O. mercator is more frequent on oil-seed products and more temperature sensitive. They enter damaged grains and feed specially on the germ. Optimum conditions for development are between 30- 350C and 70-90 percentage relative humidity.

Natural history:
Adults are 3 mm flattened narrow winged beetles but they rarely fly. Females lay their eggs loosely within the stored products. Larvae are free living and start by feeding on the embryo and the endosperm. They require 60-90 percentage humidity for optimal development, and neither species cannot develop or breed at temperatures less than 190C. All stages die in ten minutes if exposed to 550C (Howe, 1956; Halstead, 1980).

Dermestidae (Skin beetles)
Members of this family are ovoid in shape with hairy or sometimes scaly bodies. Larvae are very hairy. When stores are infested, these setae may be seriously hazardous if inhaled by workers. This family contains a number of very destructive and economically important species. One of the most serious stored product pests that belong to this family is the khapra beetle: Trogoderma granarium Everts. Apparently the only phytophagous species in the genus Trogoderma. A native of India, the Khapra beetle is now found in most parts of the world specially hot and dry areas. Adults are oval, red brown insects with a dark thorax. Adult females may lay up to 120 eggs within the stored products. Larvae are considered primary pests as they attack undamaged grains and seeds and bore into stored pulses. They are highly mobile, and in the absence of food they enter a diapause that might last for more than two years, in which they can be highly resistant to the application of pesticides or fumigation. Adults are 3-4 mm long, dark wingless beetles that do not feed. Populations of this pest build up rapidly, especially in the hot humid tropics. This species was apparently eradicated in the United States and the former Soviet Union. It also seems to be absent from East and southern Africa (Banks, 1977; Rebolledo & Arroyo, 1995; Sudesh et al., 1996 b).

 Anobiidae
Anobiids are cylindrical pubescent beetles, 1-9 mm in length. The head is usually concealed from above by the hoodlike pronotum. Most anobiids live in dry vegetable materials or bore in wood, while others are fungus feeders. About 1000 species of Anobiidae are known, most of which are found in the tropics. The following are two widespread storage pests belonging to this family.

Cigarette beetle: Lasioderma serricorne (Fabricius) is a common pest of stored cereals, cocoa beans, tobacco, ground nut, peas, beans, flours and other foodstuffs. Originally from South America, it is now found in most of the warmer parts of the world. This species is notorious for attacking a wide range of intact cereal grains, pulse seeds and food stuffs.


Natural history:
Adults can breed anywhere at optimum temperatures of around 28-320C and a relative humidity of 75 percentage. Newly hatched larvae are very active and responsible for most of the damage. Adults are small brown beetles and the only damage they cause is due to their emergence holes. This pest can be controlled if exposed to temperatures below 180C. At 550C, all stages die in two hours (see Howe, 1957; Lefkovitch & Currie, 1967).

Drug store beetle: Stegobium paniceum (Linnaeus) Another widespread pest that infests several cereals, but less common than L. serricorne in the tropics.

Lepidoptera
Lepidoptera is the second most important order of insects pests of stored products. Adults are active flyers with two pairs of scaly wings. Mouthparts of the adults are modified to suck plant nectar or other fluids and are not able to chew, while those of the larvae possess well-developed mandibles. Larvae are distinguished from beetle larvae by their pseudopods (false legs) on some of the abdominal segments. Lepidoptera larvae occur frequently in a wide range of habitats and are known for their silk-spinning activities that result in the additional loss of quality of stored products. Some species attack the product in both the field and store. Several moths are pests of the ripening crop and their larvae can be found in recently harvested stored grains. They either continue their attack for a short time in the store or form an entry point for further attack by true storage pests. The following families contain the most economically important lepidoptera post-harvest pests.

Pyralidae
Pyralidae is a large family, of which only a few species are stored product pests. Most pyralids are small and delicate moths. Members of this family exhibit a great deal of variation in appearance and habits. Larvae of all species possess glands which secrete silk with which they interlink food products as they move. This family is divided into a number of subfamilies, with the subfamily Phycitinae containing some of the most important stored grain pests. The best-known species in this subfamily are the following:

The Mediterranean flour moth: Anagasta kuehniella (Zeller).

Adults are similar to E. cautella but the body is relatively longer. A major pest of flour mills, its main habitats are flour and grout mills, corn milling plants, bakeries and any other place used for processing grains or preparing flour products. E. kuehniella occurs in most of the temperate and sub-tropical parts of the world, where average temperatures are around 200C-250C. Complete development requires about 74 days at 250C and 75 percentage relative humidity. Larvae entwine all the material on which they feed resulting in solid lumps of food particles, faeces and larval exuviae (see Jacob & Cox, 1977; Locatelli & Biglia, 1995).

Indian meal moth: Plodia interpunctella (Hübner)

This insect feeds mainly on meals and flours but can attack raisins, nuts and some pulses and whole cereals. The Indian meal moth is distributed all over the tropics and sub-tropics and in some parts of the temperate regions, specially in heated buildings. In the hot tropics, it is more abundant in cooler highland areas. Most of the damage occurs due to larval feeding on the germinal part of the grains. Damage also occurs through the contamination of foodstuff with dead larvae, frass and silk webbing.

Natural history:
Larvae feed in tubes they weave from silk secretions. Adult females stick about 200-400 eggs to the substrate or to the storage walls. Larvae develop and feed within the substrate and are sensitive to changes in temperature. The number of generations may be only two per year in Europe, but increases in the tropics to eight generations. Complete development takes about 27 days at 300C and 70 percentage relative humidity. Development ceases below 150C. All stages die at 550C in five hours (see Bell, 1975; Aitken, 1984; Locatelli & Biglia, 1995).

Gelechiidae
Gelechiidae is a large family of lepidoptera. All moths are small in size and several species are important plant pests. This family contains two serious post-harvest pests:

Angoumois grain moth: Sitotroga cerealella (Olivier)

The presence of pheromone in Angoumois grain moth, Sitotroga cerealella was first demonstrated by KEYS and MILLS (1968) using extraction of a sex attractant from female and described it as benzene.  The actual pheromone chemicals of the angoumois grain moth was reported by  VICK       et al. (1974) followed  by SU and MAHANY (1974); HAMMOUD and DESCOINS, 1978; ODINOKOV et al., 1998) as (Z, E)-7,11-hexadecadien-1-ol acetate.  This single component is one of the two major components along with Z7E11-16Ald of the pheromones of citrus leafminer moth, Phyllocnistis citrella Stainton (Lepidoptera: Phyllocnistidae) (ANDO et al., 1985).  Some of the other species utilize (Z,E)-7,11-Hexadecadienyl acetate in its chemical communication system are Uliaria rasilella (Lepidoptera:Gelechiidae), Dichomeris oceanis (Leidoptera:, Gelechiidae), Scrobipalpa sp. (Lepidoptera: Gelechiidae),    Pectinophora endema; Pink bollworm Pexicopia malvella; Hollyhock seed moth Pexicopia sp. (Lepidoptera: Gelechiidae),  Stathmopoda theoris (Lepidoptera: Stathmopodidae), Diarsia canescens (Lepidoptera: Noctuidae) (http://www.herobase.net/ database/ compound /compounds - detail- Z7E11-16A c. php).

 

In India, the sticky traps developed at the Central Food Technological Institute in Mysore, India, baited with female pheromone were compared with a commercially available Zoecon Corporation trap for monitoring the stored products pest, S. cerealella in rice stores at Mysore, India. The sticky trap was superior to the Zoecon trap but tend to collect dust and dry quickly (Karan-Singh and Majumder 1989). Apart from this information no other work was reported on this insect. In Western countries monitoring of Angoumois grain moth the delta trap is baited with rubber dispenser is used in godown. Based on the experience gained in India of working on various families/orders of insects of major crops, there are possibilities of improving the trap for efficient trapping was felt.  This has prompted us to think of using some locally available materials for use as trap  for trapping of males more efficiently.


This species is a serious primary pest that mainly attacks maize, wheat and sorghum, both in the field and in stores. A recent survey in southern Ethiopia revealed that this pest alone was responsible for 11.2 to 13.5 percentage weight loss in stored maize (Emana & Assefa, 1998). Infestation with S. cerealella starts in the field as females lay their eggs, singly or in groups, on grains. Larvae start feeding inside the grains, while still in the milk stage, and spend their entire life inside one grain. Thus, infestation is difficult to detect at this stage. Adults leave a conspicuous emergence hole at one end of the kernel. Infested grains are characterised by this circular window created by the larvae. Stored grains may be completely destroyed. Adults are active fliers, thus, they are able to infest neighbouring granaries, which is known as "cross-infestation". This pest is distributed throughout the warmer parts of the world (Africa, South and Latin America and southern Asia and Australia) (Grewal and Atwal, 1969; Boldt, 1974)

In a recent study, Nandagopal  et al., (2010) reported an efficient trap for this insect in storage. Based on the results as given above,  the recommendation for the use of  delta trap for trapping the males of S. cereallela  was found to be sufficient for monitoring of the occurrence of this species.  However, the purpose of recommending a plastic trap which proved its efficiency of significantly  higher mass trapping of the fresh or established population S. cereallela.  The return movements of the males which orient 70 º angle of return facilitated for expanding the area of trapping the males from delta trap to a simple plastic trap. During the course of observation /counting of trapped males it was noted that the entire area of the trappable area were filled with males.  This has suggested to go for expanding  the area further, which proved to increase the number of males trapped to an increase of 101%.

 

To conclude, the use of plastic tray fitted with the pheromone lure is recommended for mass trapping in the stored paddy.  Further studies on the number  of traps to be used for monitoring/ unit area, number of trap used for mass trapping/unit area, longevity of the lure (self life) under godown conditions are under way.


Potato tuberworm: Phthorimaea operculella (Zeller) = (Gnorimoschema operculella (Zeller))

This species is a cosmopolitan pest of potatoes, tomatoes and eggplants. It attacks plants mainly in the field, but continues to feed on tubers in storage. Larvae mine in the leaves and stems and later bore into the tubers. Damage can be seen on leaves as silver spots due to the tunnelling larvae, or as tunnels in the plant stem.

Natural history:
Each female lays about 150-200 eggs and larvae tunnel through leaves and stem down to the tuber where pupation may take place. In the store, eggs are laid individually on the tubers near the eyes or on sprouts. P. operculella is an important pest in traditional potato stores in North Africa (Arx et al., 1987; Lagnaoui et al., 1996. See also Haines, 1977). High infestations of up to 50 percentage of tubers can take place in Yemen due to this pest (Kroschel, 1994).


Acaridae
Flour mite: Acarus siro
Mites are widely distributed tiny arthropods. They can live and develop on various plant in the field or indoors. Mites can be found in granaries, feed mixing plants, threshing floors, stacks of hay and straw, dead organic matter, soil or plant residues. Several species are predacious on other mites or insects. Mites are easily transmitted by virtue of their tiny size which allows them to be carried with dust, winds, insects, birds or rodents. About 30 mite species are known to be associated with stored products. Family Acaridae contains some damaging species, in which Acarus siro is probably the most important and commonly encountered mite in granaries. This mite is about 0.7 mm in length with an oval body. A. siro is a widely distributed polyphagous species that can be found on almost all products of plant or animal origin. It requires relatively high humidity (70 percentage), with humidities below 11 percentage being lethal to the mite. Temperatures below -150C for 24 hours kill all stages. At 600C, all stages die in 5 minutes.

Attacked grains lose nutrients and the ability to germinate due to feeding on the germ. Crushed bodies of Acarus cause coloration in flour that reduces the products value. Under normal conditions, this mite develops according to the following pattern: egg, larva, nymph I, nymph II, and adult. Some strains of A. siro may produce hypopus under favourable conditions. Hypopus is a diapause form that can be carried by rodents or insects to other storing places. However, this species does not seem to occur in most of the tropical lowlands, though it might sometimes infest grains in cooler upland areas (see Haines, 1991).


Insect
Pheromone
Purpose
Reference
 Linnaeus
4OH :
Butan-1-ol
3-Methylbutan-1-ol
5Ald :
Pentanal
maltol :
2-Methyl-3-hydroxy-4-pyranone
4-Hydroxy-3-methoxybenzaldehyde

A
Germinara, 2008 


                                               E2  9Ald;
(E)-2-Nonenal
1-(4-Eethylphenyl)-ethanone

A
Collins, 2007 


(2S,3R)-1-Ethylpropyl 2-methyl-3-hydroxypentanoate

PM
Agg. pheromone 
Chambers,1996                  


Phillips,   1989       


Phillips,1987


(4S,5R)-5-Hydroxy-4-methylheptan-3-one
5Ald :
Pentanal
2-Methyl-3-hydroxy-4-pyranone
4-Hydroxy-3-methoxybenzaldehyde

Phillips, 1993 


(4S,5R)-5-Hydroxy-4-methylheptan-3-one 


P
Agg. pheromone 
Phillips, 1985 



PM
Agg. pheromone 
 Schmuff,  1984 

Sitophilus zeamais Motschulsky
(4S,5R)-5-Hydroxy-4-methylheptan-3-one 
PL
Hodges,1998  

2-(4-Methylcyclohex-3-enyl)-propan-2-ol


AL
Repellant
Ndungu,  1995 

(4S,5R)-5-Hydroxy-4-methylheptan-3-one 
PM
Agg. pheromone 
Walgenbach, 1987 


(4S,5R)-5-Hydroxy-4-methylheptan-3-one 

P
Agg. pheromone 
Phillips, 1985 


(4S,5R)-5-Hydroxy-4-methylheptan-3-one  
PM
Agg. pheromone 






 4,8-Dimethyldecanal
 1-Pentadecene
P
Agg Pheromone
Arnaud,  2002               J. Chem. Ecol.               28:   523 

4,8-Dimethyldecanal

PM
Agg Pheromone
Phillips,     1993                  J. Chem. Ecol. 19:723 

4,5-Dimethyldecanal
P
Agg Pheromone
Rangaswamy, 1991b 

(Z)-2-Nonenyl propionate
PF
Rangaswamy,1991a 

4,8-Dimethyldecanal

PF
Agg Pheromone
Mondal Kamesh, 1984 


4R,8S)-4,8-Dimethyldecanal


4R,8R)-4,8-Dimethyldecanal


PM
Agg Pheromone
Suzuki,  1984



4R,8R)-4,8-Dimethyldecanal


PM
Agg Pheromone
Suzuki,  1983



Tribolure ng3.2:
4,8-Dimethyldecanal
Agg Pheromone
Suzuki,  1979 


2-Ethyl-1,4-benzoquinone

toluquinone (1.0):
2-Methyl-1,4-benzoquinone

1,4-Benzoquinone

delta1-15Hy- (15.8) :
1-Pentadecene

P
Suzuki,  1978 

4,8-Dimethyldecanal
1-Pentadecene
P
Agg Pheromone
Arnaud,  2002               J. Chem. Ecol. 28: 523


Tribolure:
4,8-Dimethyldecanal
PM
Agg Pheromone
Suzuki,  1979 


2-Ethyl-1,4-benzoquinone
toluquinone (1.0):
2-Methyl-1,4-benzoquinone
1,4-Benzoquinone
delta1-15Hy- (17.6) :
1-Pentadecene
AI
Engelhardt, 1978                        24:785

1-Pentadecene

16Hy:
Hexadecane
1-Heptadecene
Heptadecadiene

P
M & F
Agg. Pheromone
Keville,  1975



1-Pentadecene
-Methyl-1,4-benzoquinone

2-Ethyl-1,4-benzoquinone

AI
M & F
Defense substance
Endt, 1971 


-Methyl-1,4-benzoquinone

2-Ethyl-1,4-benzoquinone

AI
M & F
Defense substance
 Engelhardt, 1965 

Tenebrio molitor L
 (Z)-3-Dodecenyl acetate
PM
Bryning,    2005             


2-Methyl-1,4-benzoquinone

3-Methylphenol


AI
M & F
Attygalle,1991a   


4me-9OH  30:


P  F ng
Tanaka,    1986              

Rhizopertha dominica (Fabricius)
(S)-1-Methylbutyl (E)-2-methyl-2-pentenoate

(S)-1-Methylbutyl (E)-2,4-dimethyl-2-pentenoate

P L mg
Agg.
Pheromone
Edde,    2005              Environ. Entomol.       34:             1549 

(S)-1-Methylbutyl (E)-2-methyl-2-pentenoate

 (S)-1-Methylbutyl (E)-2,4-dimethyl-2-pentenoate

P M
Agg.
Pheromone
Bashir,  2001                  


(S)-1-Methylbutyl (E)-2,4-dimethyl-2-pentenoate

(S)-1-Methylbutyl (E)-2-methyl-2-pentenoate

P L
Agg.
Pheromone
Cogburn,1984 


 (S)-1-Methylbutyl (E)-2-methyl-2-pentenoate

P L
Agg.
Pheromone
Hodges,1983 

(S)-1-Methylbutyl (E)-2-methyl-2-pentenoate

(S)-1-Methylbutyl (E)-2,4-dimethyl-2-pentenoate
P
Agg.
Pheromone
Williams HJ         1981a                   J. Chem. Ecol.          7:759



(S)-1-Methylbutyl (E)-2,4-dimethyl-2-pentenoate

P
Agg.
Pheromone
Khorramshahi, 1981      

Oryzaephilus surinamensis (L)
(E)-2-Nonenal

1-(4-Eethylphenyl)-ethanone

A
Collins, 2007 



 1-Octen-3-ol
 3-Methylbutan-1-ol

1-Octen-3-one
Octan-3-ol
8-3Kt :
Octan-3-one

2-Phenylethanol

Ethanol

K  H
Pierce,1991b 


cucujolide IV (4.4):
(Z,Z)-3,6-Dodecadien-11-olide

cucujolide IX (1.0):
(Z,Z)-3,6-Dodecadienolide

cucujolide V (2.0):
(Z,Z)-3,6-Dodecadienolide


1-Octen-3-ol
8-3Kt :
Octan-3-one

9Ald :
Nonanal

P  M
Agg.
Pheromone
White,    1989                


cucujolide IV (4.4):

(Z,Z)-3,6-(11R)-Dodecadien-11-olide
cucujolide V (2.0) :
(Z,Z)-5,8-Tetradecadien-13-olide

cucujolide IX (1.0):
(Z,Z)-3,6-Dodecadienolide

(Z)-5-Tetradecen-13-olide

P
Agg.
Pheromone
Pierce,     1987



2-Methylpropanoic acid
4Acid :
Butyric acid
2-Methylbutanoic acid
Hexanoic acid
K  H
Stubbs,      1985              


cucujolide IV (4.4):
(Z,Z)-3,6-Dodecadien-11-olide

cucujolide IX (1.0):
(Z,Z)-3,6-Dodecadienolide

cucujolide V (2.0):
(Z,Z)-5,8-Tetradecadien-13-olide

P  M
Agg.
Pheromone
Pierce,     1985 


(E)-2-Nonenal
(E,E)-2,4-Nonadienal
6Ald :
Hexanal
7Ald :
Heptanal
8Ald :
Octanal
(E)-2-Heptenal
Furan-2-carbaldehyde
3Ald :
Propanal
  Formaldehyde
A
M |& |F
Mikolajczak, 1984 


 Propanal

F
O'Donnell, 1983 

Oryzaephilus mercator (Fauvel)
1-Octen-3-ol
3-Methylbutan-1-ol
1-Octen-3-one
Octan-3-ol
Octan-3-one
K  H
Pierce, 1991b   


 (Z)-3-(11R)-Dodecen-11-olide
 (Z,Z)-3,6-(11R)-Dodecadien-11-olide
(Z)-5-Tetradecen-13-olide
P
Agg.
Pheromone
Pierce,     1987 


(Z)-3-(11R)-Dodecen-11-olide
(Z,Z)-3,6-(11R)-Dodecadien-11-olide
P
Agg.
Pheromone


(Z,Z)-3,6-(11R)-Dodecadien-11-olide
(Z)-3-(11R)-Dodecen-11-olide
P M
Agg.
Pheromone
Pierce,     1985 

 Trogoderma granarium Everts
(R)-(Z)-14-Methyl-8-hexadecenal
(R)-(E)-14-Methyl-8-hexadecenal
P  F
Levinson HZ        1980 


(S)-(Z)-14-Methyl-8-hexadecenal
(S)-(E)-14-Methyl-8-hexadecenal
P  F
Rossi,1979 


S)-(Z)-14-Methyl-8-hexadecenal
(S)-(E)-14-Methyl-8-hexadecenal
P  F
Rossi,1978a



(E)-14-Methyl-8-hexadecenal
14-Methyl-8-hexadecen-1-ol
hexadecenoate (o.2):
Methyl 14-methyl-8-hexadecenoate
Methyl7-hexadecenoate
5-Ethyldihydro-2(3H)-furanone
E-trogodermal (0.5):
E)-14-Methyl-8-hexadecenal
caproic acid (2.98) :
Hexanoic acid
P F   ng

(Z)-14-Methyl-8-hexadecenal
(E)-14-Methyl-8-hexadecenal

P  F
Cross  et al., 1976          


Hexadecanoic acid
(Z)-9-Octadecenoic acid
(Z,Z)-9,12-Octadecadienoic acid
(Z)-9-Hexadecenoic acid
Heptadecanoic acid
Octadecanoic acid
A
Cohen   et al., 1971       

(Z)-9-Octadecenoic acid methyl ester
Ethyl hexadecanoate
Octadecanoic acid ethyl ester
(Z)-9-Octadecenoic acid ethyl ester
(Z,Z)-9,12-Octadecadienoic acid ethyl
P  M & F 
Agg. Pheromone
Ikan  et al., 1969  

Lasioderma serricorne (Fabricius)
S)-4-(Prop-1-en-2-yl)-cyclohex-1-enecarbaldehyde
S-limonene (14.42):
(S)-1-Methyl-4-(1-methylethenyl)-cyclohexene
E-beta-farnesene (11.62):
(E)-7,11-Dimethyl-3-methylene-1,6,10-dodecatriene
1R-(1R*,4E,9S*)-4,11,11-Trimethyl-8-methylenebicyclo[7.2.0]undec-4-ene
Al  Repellant
Hori,   2004b                  


2-Hydroxy-4-isopropyl-2,4,6-cycloheptatrien-1-one
Al  Repellant
Hori ,  2004a 

(2S,3R,1'S)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6(1-methyl-2-oxobutyl)-4H-pyran-4-onebeta-serricorone:
(2S,3R,1'R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6(1-methyl-2-oxobutyl)-4H-pyran-4-one
P F
Marking pheromone
Imai , 1990b 


(4S,6S,7S)-7-Hydroxy-4,6-dimethylnonan-3-one

(2S,3S)-2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran
P F



(4S,6S,7S)-7-Hydroxy-4,6-dimethylnonan-3-one
(2S,3S)-2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran
4,6-Dimethylnonan-3,7-dione
4,6-Dimethylnonan-3,7-diol
7-Hydroxy-4,6-dimethyl-4-nonen-3-one
(2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-oxobutyl)-4H-pyran-4-one
(2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one
P F

Chuman ,         1985      


(4S,6S,7S)-7-Hydroxy-4,6-dimethylnonan-3-one
P F

Mochizuki   1984 


 (2S,3S)-2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran
7-Hydroxy-4,6-dimethylnonan-3-one
P F

Levinson HZ        1981 


7-Hydroxy-4,6-dimethylnonan-3-one
P F

Chuman et al., 1979 
Stegobium paniceum (Linnaeus)
(2S,3R,1'S)-2,3-Dihydro-2,3,5-trimethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one

Kodama,  1987b



2,3-Dihydro-2,3,5-trimethyl-6-(1-methyl-2-oxobutyl)-4H-pyran-4-one

 Kodama,1987a 


2,3-Dihydro-2,3,5-trimethyl-6-(1-methyl-2-oxobutyl)-4H-pyran-4-one

Kuwahara,1978 
                  

(E)-3,7,11,15-Tetramethyl-2-hexadecen-1-ol
5-Hexyl-dihydrofuran-2(3H)-one
5-Heptyl-dihydrofuran-2(3H)-one
P M
Phelan,   1986 


Z9E12-14Ac  (3.0):
(Z,E)-9,12-Tetradecadienyl acetate
Z9E12-14OH (0.3):
(Z,E)-9,12-Tetradecadien-1-ol
P F ng
Krasnoff,1984 


 (Z,E)-9,12-Tetradecadienyl acetate
Al
Soldan,  1983 


Z9E12-14Ac  (7.6):
(Z,E)-9,12-Tetradecadienyl acetate
Z9E12-14OH (2.0):
(Z,E)-9,12-Tetradecadien-1-ol
P F ng
Kuwahara, 1973 

(Z,E)-9,12-Tetradecadienyl acetate
P F
 Brady,    1971b           

 Ephestia kuehniella (=Anagasta kuehniella) (Zeller).
Phenylmethanol
9Ald :
Nonanal
2-Phenylacetaldehyde
A
Olsson,2006 


3me-4OH (1.0):
3-Methylbutan-1-ol
acetic acid (1.0):
Acetic acid
Al
 Toth,2002a 

Z9E12-14Ac  (5.6):
(Z,E)-9,12-Tetradecadienyl acetate
Z9E12-14OH (1.2):

(Z,E)-9,12-Tetradecadien-1-ol
P F  ng
Kuwahara,1973 


(Z,E)-9,12-Tetradecadienyl acetate
P F 
Kuwahara, 1971b 
Plodia interpunctella (Hübner)
(Z,E)-9,12-tetradecadien-1-ol: acetate:

F P
Sower et al., 1974

Phthorimaea operculella (Zeller) (=Gnorimoschema operculella (Zeller))
E4Z7-13Ac (4.0):
(E,Z)-4,7-Tridecadienyl acetate

E4Z7Z10-13Ac (6.0):
(E,Z,Z)-4,7,10-Tridecatrienyl acetate

P  L
Ono et al.,           1997

E4Z7-13Ac (8.6):
(E,Z)-4,7-Tridecadienyl acetate
E4Z7Z10-13Ac (3.9):
(E,Z,Z)-4,7,10-Tridecatrienyl acetate

P F ng
Ono  et al.,          1997                     


E4Z7-13Ac (2.7):
(E,Z)-4,7-Tridecadienyl acetate
E4Z7Z10-13Ac (1.4):
(E,Z,Z)-4,7,10-Tridecatrienyl acetate

P F ng
Ono   et al., 1997           


E4Z7-13Ac (3.5):
(E,Z)-4,7-Tridecadienyl acetate
E4Z7Z10-13Ac (3.7):
(E,Z,Z)-4,7,10-Tridecatrienyl acetate
P F ng

(E,Z)-4,7-Tridecadienyl acetate

P F
Raman,  1982

E4Z7-13Ac (9.0):
(E,Z)-4,7-Tridecadienyl acetate
E4Z7Z10-13Ac (1.0):
(E,Z,Z)-4,7,10-Tridecatrienyl acetate
P L

E4Z7-13Ac (4.0):
(E,Z)-4,7-Tridecadienyl acetate
E4Z7Z10-13Ac (1.0):

P L

(E,Z)-4,7-Tridecadienyl acetate
(E,Z,Z)-4,7,10-Tridecatrienyl acetate
P F

(E,Z)-4,7-Tridecadienyl acetate
P

E4Z7Z10-13Ac :(E,Z,Z)-4,7,10-Tridecatrienyl acetate
P F

(E,Z,Z)-4,7,10-Tridecatrienyl acetate
P F

(Z,Z)-7,11-Tridecadienyl acetate
P

(Z)-5-Dodecenyl acetate
A L
Steck,  1979c 

Flour mite: Acarus siro L.
guanine (2-amino-6-hydroxypurine):


Assembly pheromone
Levinson et al,.1991

Ammonia:
Kairomone  
For both sex
Levinson et al,.1991

Tridecane
(Z)-3,7-Dimethyl-2,6-octadienal
(E)-3,7-Dimethyl-2,6-octadienal

P
Tuma et al., 1990
(Z)-3-Methyl-2-heptenoic acid
P  F
Cork,  1991b 

2,6-Dimethyl-1,8-octanedioic acid

(E)-3,7-Dimethyl-2-octene-1,8-dioic acid
P
Shimomura et al., 2010
(Z,E)-7-Ethyl-3,11-dimethyl-2,6,10-dodecatrienal
(E,E)-7-Ethyl-3,11-dimethyl-2,6,10-dodecatrienal
P F
2008 


Hexane
3-2Kt :
Propan-2-one
Ethanol
Methanol
A
Khan, 1998  


3-Methylpentacosane
11-Methylheptacosane
3-Methylheptacosane
11-Methylnonacosane
13-Methylnonacosane
11,15-Dimethylnonacosane
13-Methylhentriacontane
9,13-Dimethylhentriacontane
11,15-Dimethyltritriacontane
callosobruchusic acid :
(E)-3,7-Dimethyl-2-octene-1,8-dioic acid
P F
Tanaka, 1981                 

Callosobruchus maculatus
 Fabricius
azelaic acid (2.5):
Nonanedioic acid
2,6-Dimethyl-1,8-octanedioic acid
P F ng
Contact pheromone
Nojima,  2007                


3-Methyleneheptanoic acid
(Z)-3-Methyl-3-heptenoic acid
(E)-3-Methyl-3-heptenoic acid
(Z)-3-Methyl-2-heptenoic acid
(E)-3-Methyl-2-heptenoic acid
P F
(Z,E)-7,11-Hexadecadienyl acetate
(Z,E)-7,11-Hexadecadienal
P L
Ando et al., 1985

Hexadecadienyl acetate
P F
Vick et al., 1974

P L
Keys et al., 1968

A-Atractant, Al-Allomone, K-Kairomone, P-Pheromone, L-Larvae, F-Female,
H-   Host, M-Male, M&F-male and Female, ng-nanogram

          List of various types of traps practiced in trapping the storage insects

Name of insect
Type of trap
Result
Reference

Tobacco beetle, Lasioderma serricorne F.
Traps
The pheromone capsule dispenser captured the most adults and Serrico lure captured the least
Trematerra
et al.,1978

Rhyzopertha dominica
Window trap
The window trap proved  the most effective for monitoring this pest in wheat grains
Singh and Sinha,1998

Sweet potato weevil,
Cylas formicarius (Fabricius)
Root traps
No weevils were found upon re-examination of root traps and dissection of wild host plants carried out in September, 1996
Setokuchi, 2001

Pheromone traps
The sticky trap was superior to the Zoecon trap but tended to collect dust on the sticky surface.
Lal, 1990


Pheromone traps
It was concluded that SPW had been completely eradicated from Muroto City
Komi, 2000

Sweet potato weevil,
Cylas formicarius elegantulus
Plastic funnel traps
Plastic funnel traps caught significantly more weevils (90-97% of the total catch) and were significantly more efficient (85%  efficiency)
Jansson et al., 1989

Stored products pests
Ready-made insect traps
In situations where slight inaccuracy in readings can be tolerated, the efficiency of the sensors as insect traps could be increased by loading them with the most attractive filler, treated with an insecticide
Hodges, 1983

Cucujid and tenebrionid beetles
Plastic probe traps, corrugated cardboard traps and weighing boat traps
For each target species, pheromone, trap type, and habitat development of optimal monitoring programmes should be  independent
Javer et al., 1990

Khapra beetle
Wall-mounted trap
Traps mounted on a wall did not work unless flaps were deployed
Barak,1989

Storage insect pests
Corrugated paper insect trap, plastic probe trap
 More insect pests were trapped with corrugated paper traps than with the other devices tested in grain storage
Ayertey,
1989

Adult beetles (Coleoptera)
Probe trap, the Storgard WB Probe II (WB II) and Grain Guard (GG)
Insects in shelled maize with WB II traps and in stored barley with GG traps  catches with   84% variation
Subramanyam et al.,1993

Sitophilus granarius
Pitfall beaker, insect probe and WBII probe traps
Correlation between trap catch increase and increase in grain temperature
Wakefield and Cogan,1999

Oryzaephilus surinamensis, Sitophilus granarius
and
Cryptolestes ferrugineus
Pitfall and probe traps
The possibility of use of nuclear magnetic resonance to detect those insects and their immature stages hidden within grains was discussed
Pinniger et al.,1986

Tribolium castaneum
Pitfall and probe trap
The behavioural responses of the beetles to the pheromone in still and moving air were similar
Obeng-Ofori and Coaker,1990

Flour beetles,
Tribolium confusum
and
T. castaneum
Pitfall traps
Pitfall traps were efficient
Fisher et al.,1993

Stored product beetles
Pitfall trap
Pitfall trap was efficient
Obeng-Ofori,1993
Flour moth,
Ephestia kuehniella

A strain of the parasitoid Trichogramma evanescens and the predator Blattisocius tarsalis have been selected for further investigation
Hansen et al., 1998
Potato tuberworm, Phthorimaea operculella
Dry and a water trap
Pheromone traps were efficient in trapping the insects
Karan–Singh et al., 1989
Water pan trap
Water pan trap was efficient in trapping the males of the potato tuber moth
Tomio and  Orita, 1984
Grain moth,
Sitotroga cerealella Olivier
Sticky traps, Zoecon trap
All the traps baited with synthetic pheromones caught more males than those baited with virgin females, with the most moths (837.33/trap) being caught with a 1:1 ratio of the components


Tray trap

increase the number of males trapped to an increase of 101%.

Nandagopal
et  a l.,2010
Cylas formicarius,
Ips typographus,
Tomicus piniperda,
Pityogenes chalcographus, Scolytus triarmatus
Pheromone traps
The sticky trap was superior to the Zoecon trap but tended to collect dust on the sticky surface.
Lal, 190


References

Ando, T., Taguchi, K., Uchiyama, M., Ujiye, T., and Kuroko, H. 1985b. (7Z,11Z)-7,11-hexadecadienal: sex attractant of the citrus leafminer moth, Phyllocnistis citrella Stainton (Lepidoptera, Phyllocnistidae). Agric. Biol. Chem. 49: 3633-3635.
Arnaud, L., Lognay, G., Verscheure, M., Leenaers, L., Gaspar, C., and Haubruge, E. 2002. Is dimethyldecanal a common aggregation pheromone of Tribolium flour beetles? J. Chem. Ecol. 28: 523-532.
Attygalle, A.B., Blankespoor, C.L., Meinwald, J., and Eisner, T. 1991a. Defensive secretion of Tenebrio molitor (Coleoptera: Tenebrionidae). J. Chem. Ecol. 17: 805-809.
Babu, A.,  A. Hern and S. Dorn 2003. Sources of semiochemicals mediating host finding in Callosobruchus chinensis (Coleoptera: Bruchidae). Bulletin of Entomological Research, 93: 187-192.
Bacon, O.G., Seiber, J.N., and Kennedy, G.G. 1976. Evaluation of survey trapping techniques for potato tuberworm moths with chemical baited traps. J. Econ. Entomol. 69: 569-572.
 Bashir, T., Birkinshaw, L.A., Farman, D., Hall, D.R., and Hodges, R.J. 2001. Pheromone release by Rhyzopertha dominica (F) (Coleoptera: Bostrichidae) in the laboratory: daily rhythm, inter-male variation and association with body weight and/or boring activity. J. Stored Prod. Res. 39: 159-169.
Brady, U.E., and Nordlund, D.A. 1971b. Cis-9,trans-12 tetradecadien-1-yl acetate in the female tobacco moth Ephestia elutella (Hübner) and evidence for an additional component of the sex pheromone. Life Sci. 10: 797-801.
Bryning, G.P., Chambers, J., and Wakefield, M.E. 2005. Identification of a sex pheromone from male yellow mealworm beetles, Tenebrio molitor. J. Chem. Ecol. 31: 2721-2730.
 Chambers, J., Van Wyk, C.B., White, P.R., Gerrard, C.M., and Mori, K. 1996. Grain weevil, Sitophilus granarius (L.): antennal and behavioral responses to male-produced volatiles. J. Chem. Ecol. 22: 1639-1654.
Chuman, T., Kohno, M., Kato, K., and Noguchi, M. 1979. 4,6-dimethyl-7-hydroxy-nonan-3-one, a sex pheromone of the cigarette beetle (Lasioderma serricorne F.). Tetrahedron Lett. 25: 2361-2364.
Chuman, T., Mochizuki, K., Mori, M., Kohno, M., Kato, K., and Noguchi, M. 1985. Lasioderma chemistry. Sex pheromone of cigarette beetle (Lasioderma serricorne F.). J. Chem. Ecol. 11:417-434.
Cogburn, R.R., Burkholder, W.E., and Williams, H.J. 1984. Field tests with the aggregation pheromone of the lesser grain borer (Coleoptera: Bostrichidae). Environ. Entomol. 13: 162-166.
Cohen, E., Ikan, R., and Sulov, A. 1971. Response of the females of Trogoderma granarium (Coleoptera, Dermestidae) to cuticular free fatty acids. Entomol. Exp. Appl. 14: 315-318.
Collins, L.E., Bryning, G.P., Wakefield, M.E., Chambers, J., and Cox, P.D. 2007. Progress towards a multi-species lure: identification of components of food volatiles as attractants for three storage beetles. J. Stored Prod. Res. 43: 53-63.
Collins, L.E., Bryning, G.P., Wakefield, M.E., Chambers, J., and Cox, P.D. 2007. Progress towards a multi-species lure: identification of components of food volatiles as attractants for three storage beetles. J. Stored Prod. Res. 43: 53-63.  
Cork, A., Hall, D.R., Blaney, W.M., and Simmonds, M.S.J. 1991b. Identification of a component of the female sex pheromone of Callosobruchus analis (Coleoptera: Bruchidae). Tetrahedron Lett. 32: 129-132.
Cross, J.H., Byler, R.C., Cassidy, R.F., Jr., Silverstein, R.M., Greenblatt, R.E., Burkholder, W.E., Levinson, A.R., and Levinson, H.Z. 1976. Porapak-Q collection of pheromone components and isolation of (Z)- and (E)-14-methyl-8-hexadecenal, sex pheromone components, from the females of four species of Trogoderma (Coleoptera: Dermestidae). J. Chem. Ecol. 2: 457-468.
Edde, P.A., Phillips, T.W., and Toews, M.D. 2005. Responses of Rhyzopertha dominica (Coleoptera: Bostrichidae) to its aggregation pheromones as influenced by trap design, trap height, and habitat. Environ. Entomol. 34: 1549-1557.
Endt, D.W., and Wheeler, J.W. 1971. 1-pentadecene production in Tribolium confusum. Science. 172: 60-61.
Engelhardt, M., Rapoport, H., and Sokoloff, A. 1965. Odorous secretion of normal and mutant Tribolium confusum. Science. 150: 632-633.
Fouda, H.G., Seiber, J.N., and Bacon, O.G. 1975. A potent sex attractant for the potato tuberworm moth. J. Econ. Entomol. 68: 423-427.
Germinara, C.S., De Cristofaro, A.,and Rotundo, G. 2008. Behavioral responses of adult Sitophilus granarius to individual cereal volatiles. J. Chem. Ecol. 34:523-529.
Greenblatt, R.E., Burkholder, W.E., Cross, J.H., Byler, R.C., and Silverstein, R.M. 1976. Chemical communication in the mating behavior of Trogoderma glabrum (Herbst) (Coleoptera: Dermestidae). J. Chem. Ecol. 2: 285-297.
Greenblatt, R.E., Burkholder, W.E., Cross, J.H., Cassidy, R.F., Jr., Silverstein, R.M., Levinson, A.R., and Levinson, H.Z. 1977. Chemical basis for interspecific responses to sex pheromones of Trogoderma species (Coleoptera: Dermestidae). J. Chem. Ecol. 3: 337-347.
H. Z. Levinson, A. R. Levinson and K. Müller.1991. Functional adaptation of two nitrogenous waste products in evoking attraction and aggregation of flour mites (Acarus siro L.) Anzeiger für Schädlingskunde. 64(3): 55-60.
Hodges, R.J., Hall, D.R., Golob, P., and Meik, J. 1983. Responses of Prostephanus truncatus to components of the aggregation pheromone of Rhyzopertha dominica in the laboratory and field. Entomol. Exp. Appl. 34: 266-272.
 Hodges, R.J., Hall, D.R., Mbugua, J.N., and Likhayo, P.W. 1998. The responses of Prostephanus truncatus (Coleoptera: Bostrichidae) and Sitophilus zeamais (Coleoptera: Curculionidae) to pheromone and synthetic maize volatiles as lures in crevice or flight traps. Bull. Entomol. Res. 88: 131-139.
Hori, M. 2004a. Repellency of hinokitiol against the cigarette beetle, Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae). Appl. Entomol. Zool. 39: 521-526.
Hori, M. 2004b. Repellency of shiso oil components against the cigarette beetle, Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae). Appl. Entomol. Zool. 39: 357-362.
Ikan, R., Bergmann, E.D., Yinon, U., and Sulov, A. 1969. Identification, synthesis and biological activity of an "assembling scent" from the beetle Trogoderma granarium. Nature. 223: 317.
Imai, T., Kodama, H., Chuman, T., and Kohno, M. 1990b. Female-produced oviposition deterrents of the cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae). J. Chem. Ecol. 16: 1237-1247.
Keville, R., and Kannowski, P.B. 1975. Sexual excitation by pheromones of the confused flour beetle. J. Insect Physiol. 21: 81-84.
Keys, R.E., and Mills, R.B. 1968. Demonstration and extraction of a sex attractant from female angoumois grain moths. J. Econ. Entomol. 61: 46-49.
Khan, M., and Shahjahan, R.M. 1998. Evaluation of eucalyptus leaf extracts as repellent for adult Sitophilus oryzae (L.) (Col.: Curculionidae) and attractant for Callosobruchus chinensis (L.) (Col.: Bruchidae). Bangl. J. Entomol. 8: 81-88.
Khorramshahi, A., and Burkholder, W.E. 1981. Behavior of the lesser grain borer Rhyzopertha dominica (Coleoptera: Bostrichidae) male produced aggregation pheromone attracts both sexes. J. Chem. Ecol. 7: 33-38.
Kodama, H., Mochizuki, K., Kohno, M., Ohnishi, A., and Kuwahara, Y. 1987a. Inhibition of male response of drugstore beetles to stegobinone by its isomer. J. Chem. Ecol. 13: 1859-1869.
Kodama, H., Ono, M., Kohno, M., and Ohnishi, A. 1987b. Stegobiol, a new sex pheromone component of drugstore beetle (Stegobium paniceum L). J. Chem. Ecol. 13: 1871-1879.
Krasnoff, S.B., Vick, K.W., and Coffelt, J.A. 1984. (Z,E)-9,12-Tetradecadien-1-ol: a component of the sex pheromone of Ephestia elutella (Hübner) (Lepidoptera: Pyralidae). Environ. Entomol. 13: 765-767.
Kuwahara, Y., and Casida, J.E. 1973. Quantitative analysis of the sex pheromone of several phycitid moths by electron-capture gas chromatography. Agric. Biol. Chem. 37: 681-684.
Kuwahara, Y., Fukami, H., Howard, R., Ishii, S., Matsumura, F., and Burkholder, W.E. 1978. Chemical studies on the Anobiidae: sex pheromone of the drugstore beetle, Stegobium paniceum (L.) (Coleoptera). Tetrahedron. 34: 1769-1774.
Kuwahara, Y., Hara, H., Ishii, S., and Fukami, H. 1971b. The sex pheromone of the Mediterranean flour moth. Agric. Biol. Chem. 35: 447-448.
 Levinson H. Z., A. R. Levinson and K. Müller.1991. Functional adaptation of two nitrogenous waste products in evoking attraction and aggregation of flour mites (Acarus siro L.). Anzeiger für Schädlingskunde 64(3): 55-60.
 Levinson, H.Z., and Mori, K. 1980. The pheromone activity of chiral isomers of trogodermal for male khapra beetles. Naturwissenschaften. 67: 148-149.
Levinson, H.Z., Levinson, A.R., Francke, W., Mackenroth, W., and Heemann, V. 1981. The pheromone activity of anhydroserricornin and serricornin for male cigarette beetles (Lasioderma serricorne F.). Naturwissenschaften. 68: 148-149.
Markaria, H., Florentine, G.J., and Pratt, J.J., Jr. 1978. Quinone production of some species of Tribolium. J. Insect Physiol. 24: 785-790.
 Mikolajczak, K.L., Zilkowski, B.W., Smith, C.R., Jr., and Burkholder, W.E. 1984. Volatile food attractants for Oryzaephilus surinamensis (L.) from oats. J. Chem. Ecol. 10: 301-309.
Mochizuki, K., Chuman, T, Mori, M., Kohno, M., and Kato, K. 1984. Activity of stereoisomers of serricornin, sex pheromone of the cigarette beetle (Lasioderma serricorne F.). Agric. Biol. Chem. 48: 2833-2834.
Mochizuki, K., Mori, M., Chuman, T, Kohno, M., Ohnishi, A., Watanabe, H., and Mori, K. 1986. Reinvestigation of anhydroserricornin, (2S,3S)-2,6-diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran, as a sex pheromone component for male cigarette beetle. J. Chem. Ecol. 12: 179-186.
Mondal, K.A.M.S.H., and Port, G.R. 1984. Response of Tribolium castaneum larvae to synthetic aggregation pheromone. Entomol. Exp. Appl. 36: 43-46.
 Ndungu, M., Lwande, W., Hassanali, A., Moreka, L., and Chhabra, S.C. 1995. Cleome monophylla essential oil and its constituents as tick (Rhipicephalus appendiculatus) and maize weevil (Sitophilus zeamais) repellents. Entomol. Exp. Appl. 76: 217-222.
Nojima, S., Shimomura, K., Honda, H., Yamamoto, I., and Ohsawa, K. 2007. Contact sex pheromone components of the cowpea weevil, Callosobruchus maculatus. J. Chem. Ecol. 33: 923-933.
 O'Donnell, M.J., Chambers, J., and McFarland, S.M. 1983. Attractancy to Oryzaephilus surinamensis (L.), saw-toothed grain beetle, of extracts of carobs, some triglycerides and related compounds. J. Chem. Ecol. 9: 357-374.
Oehlschlager, A.C., King, G.G.S., Pierce, H.D., Jr., Pierce, A.M., Slessor, K.N., Millar, J.G., and Borden, J.H. 1987. Chirality of macrolide pheromones of grain beetles in the genera Oryzaephilus and Cryptolestes and its implications for species specificity. J. Chem. Ecol. 13: 1543-1554.
Olsson, C.P.-O., Anderbrant, O., and Löfstedt, C. 2006. Attraction and oviposition of Ephestia kuehniella induced by volatiles identified from chocolate products. Entomol. Exp. Appl. 119: 137-144.
Ono, T., Charlton, R.E., and Cardé, R.T. 1990. Variability in pheromone composition and periodicity of pheromone titer in potato tuberworm moth, Phthorimaea operculella (Lepidoptera: Gelechiidae). J. Chem. Ecol. 16: 531-542.
Ono, T., Chouvalitwongporn, P., and Saito, T. 1997. Comparison of the sex pheromone system between Japanese and Thai populations of the potato tuberworm moth, Phthorimaea operculella (Lepidoptera: Gelechiidae). Appl. Entomol. Zool. 32: 514-517.
Persoons, C.J., Voerman, S., Verwiel, P.E.J., Nooijen, P.J.F., Nooijen, W.J., Ritter, F.J., and Minks, A.K. 1976a. Sex pheromone of the potato tuberworm moth, Phthorimaea operculella: isolation and identification. Med. Fac. Landbouww. Rijksuniv. Gent. 41: 945-948.
Persoons, C.J., Voerman, S., Verwiel, P.E.J., Ritter, F.J., Nooyenand, W.J., and Minks, A.K. 1976b. Sex pheromone of the potato tuberworm moth, Phthorimaea operculella: isolation, identification and field evaluation. Entomol. Exp. Appl. 20: 289-300.
 Phelan, P.L., Silk, P.J., Northcott, C.J., Tan, S.H., and Baker, T.C. 1986. Chemical identification and behavioral charactrization of male wing pheromone of Ephestia elutella (Pyralidae). J. Chem. Ecol. 12: 135-146.
 Phillips, J.K., Chong, J.M., Andersen, J.F., and Burkholder, W.E. 1989. Determination of the enantiomeric composition of (R*,S*)-1-ethylpropyl 2-methyl-3-hydroxypentanoate, the male-produced aggregation pheromone of Sitophilus granarius. Entomol. Exp. Appl. 51: 149-153.
 Phillips, J.K., Miller, S.P.F., Andersen, J.F., Fales, H.M., and Burkholder, W.E. 1987. The chemical identification of the granary weevil aggregation pheromone. Tetrahedron Lett. 28: 6145-6146.  
 Phillips, J.K., Walgenbach, C.A., Klein, J.A., Burkholder, W.E., Schmuff, N.R., and Fales, H.M. 1985. (R*,S*)-5-hydroxy-4-methyl-3-heptanone: male-produced aggregation pheromone of Sitophilus oryzae (L.) and S. zeamais Motsch. J. Chem. Ecol. 11:1263-1274.
 Phillips, T.W., Jiang, X.-L., Burkholder, W.E., Phillips, J.K., and Tran, H.Q. 1993. Behavioral responses to food volatiles by two species of stored-product Coleoptera, Sitophilus oryzae (Curculionidae) and Tribolium castaneum (Tenebrionidae). J. Chem. Ecol. 19: 723-734.
Phillips, T.W., Phillips, J.K., Webster, F.X., Tang, R., and Burkholder, W.E. 1996. Identification of sex pheromones from cowpea weevil, Callosobruchus maculatus, and related studies with C. analis (Coleoptera: Bruchidae). J. Chem. Ecol. 22: 2233-2249.
Pierce, A.M., Pierce, H.D., Jr., Borden, J.H., and Oehlschlager, A.C. 1991b. Fungal volatiles: semiochemicals for stored-product beetles (Coleoptera: Cucujidae). J. Chem. Ecol. 17: 581-597.
Pierce, A.M., Pierce, H.D., Jr., Oehlschlager, A.C., and Borden, J.H. 1985. Macrolide aggregation pheromones in Oryzaephilus surinamensis and Oryzaephilus mercator (Coleoptera: Cucujidae). J. Agric. Food Chem. 33: 848-852.
 Pierce, A.M., Pierce, H.D., Jr., Oehlschlager, A.C., Czyzewska, E., and Borden, J.H. 1987. Influence of pheromone chirality on response by Oryzaephilus surinamensis and Oryzaephilus mercator (Coleoptera: Cucujidae). J. Chem. Ecol. 13: 1525-1542.  
Raman, K.V. 1982. Field trials with the sex pheromone of the potato tuberworm. Environ. Entomol. 11: 367-370.
Raman, K.V. 1982. Field trials with the sex pheromone of the potato tuberworm. Environ. Entomol. 11:367-370.   Steck, W.F., Chisholm, M.D., Bailey, B.K., and Underhill, E.W. 1979c. Moth sex attractants found by systematic field testing of 3-component acetate-aldehyde candidate lures. Can. Entomol. 111: 1263-1269.
Rangaswamy, J.R., and Sasikala, V.B. 1991a. Sex pheromone properties of Z-2-nonenyl propionate isolated from virgin females of red flour beetle Tribolium castaneum (Coleoptera). Indian J. Exp. Biol. 29: 263-269.
Rangaswamy, J.R., and Sasikala, V.B. 1991b. Aggregation pheromone activity of compounds isolated from male red flour beetle Tribolium castaneum (Coleoptera: Tenebrionidae). Indian J. Exp. Biol. 29: 52-55.
Roelofs, W.L., Kochansky, J.P., Cardé, R.T., Kennedy, G.G., Henrick, C.A., Labovitz, J.N., and Corbin, V.L. 1975c. Sex pheromone of the potato tuberworm moth, Phthorimaea operculella. Life Sci. 17: 699-706.
 Rossi, R., and Niccoli, A. 1978a. Relationship between chirality and biological activity. Biological response of Trogoderma granarium to optically active synthetic sex attractants. Naturwissenschaften. 65: 259-260.
Rossi, R., Salvadori, P.A., Carpita, A., and Nicoli, A. 1979. Chirality influences the biological activity of the sex pheromones of the khapra beetle. Naturwissenschaften. 66: 211.
 Schmuff, N.R., Philips, J.K., Burkholder, W.E., Fales, H.M., Chen, C.-W., Roller, P.P., and Ma, M. 1984. The chemical identification of the rice weevil and maize weevil aggregation pheromone. Tetrahedron Lett. 25: 1533-1534.
Shimomura, K., Akasaka, K., Yajima, A., Mimura, T., Yajima, S., and Ohsawa, K. 2010. Contact sex pheromone components of the seed beetle, Callosobruchus analis (F.). J. Chem. Ecol. 36: 955-965.  
Shimomura. K., Nojima, S., Yajima, S., and Ohsawa, K. 2008. Homofarnesals: female sex attractant pheromone components of the southern cowpea weevil, Callosobruchus chinensis. J. Chem. Ecol. 34: 467-477.
Shu, S.Q., Mbata, G.N., Cork, A., and Ramaswamy, S.B. 1999. Sex pheromone of Callosobruchus subinnotatus. J. Chem. Ecol. 25: 2715-2727.
 Soldan, T., and Spitzer, K. 1983. Some moths recorded at sex pheromone traps in Mitidja, Algeria (Lepidoptera: Tortricidae, Pyralidae, Noctuidae). Acta Entomol. Bohemoslov. 80: 395-398.

Sower, L. L.; Vick, K. W.; Tumlinson, J. H. 1974. (Z,E)-9,12-Tetradecadien-1-ol: A Chemical Released by Female Plodia interpunctella That Inhibits the Sex Pheromone Response of Male Cadra cautella. Environmental Entomology, 3(1): 120-122.

Steck, W.F., Chisholm, M.D., Bailey, B.K., and Underhill, E.W. 1979c. Moth sex attractants found by systematic field testing of 3-component acetate-aldehyde candidate lures. Can. Entomol. 111:1263-1269.
Stubbs, M.R., Chambers, J., Schofield, S.B., and Wilkins, J.P.G. 1985. Attractancy to Oryzaephilus surinamensis (L.) of volatile materials isolated from vacuum distillate of heat-treated carobs. J. Chem. Ecol. 11:565-581.
Suzuki, T., and Mori, K. 1983. (4R,8R)-(-)-4,8-dimethyldecanal: the natural aggregation pheromone of the red flour beetle, Tribolium castaneum (Coleoptera: Tenebrionidae). Entomol. Exp. Appl. 18:134-136.
Suzuki, T., and Sugawara, R. 1979. Isolation of an aggregation pheromone from the red flour beetles, Tribolium castaneum and T. confusum (Coleoptera: Tenebrionidae). Appl. Entomol. Zool. 14:228-230.
Suzuki, T., Nakakita, H., and Kuwahara, Y. 1984. Biological activities of the analogs of the aggregation pheromone of Tribolium castaneum (Coleoptera: Tenebrionidae). Appl. Entomol. Zool. 19:15-20.
Suzuki, T., Nakakita, H., and Kuwahara, Y. 1987. Aggregation pheromone of Tribolium freemani Hinton (Coloeoptea: Tenebrionidae) I. Identification of aggregation pheromone. Appl. Entomol. Zool. 22:340-347.
Tanaka, K., Ohsawa, K., Honda, H., and Yamamoto, I. 1981. Copulation release pheromone, erectin, from the azuki bean weevil (Callosobruchus chinensis L.). J. Pestic. Sci. 6:75-82.
 Tanaka, Y., Honda, H., Ohsawa, K., and Yamamoto, I. 1986. A sex attractant of the yellow mealworm, Tenebrio molitor L., and its role in the mating behavior. J. Pesticide Sci. 11:49-55.
Tóth, M., Bellas, T.E. and Rothschild, G.H.L. 1984. Role of pheromone components in evoking behavioral responses from male potato tuberworm moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae). J. Chem. Ecol. 10:271-280.
Tóth, M., Repasi, V., and Szöcs, G. 2002a. Chemical attractants for females of pest pyralids and phycitids (Lepidoptera: Pyralidae, Phycitidae). Acta Phytopathol. Entomol. Hung. 37:375-384.
Tuma, D., Sinha, R.N., Muir, W.E., and Abramson, D. 1990. Odor volatiles associated with mite-infested bin-stored wheat. J. Chem. Ecol. 16:713-724.
Vick, K.W., Su, H.C.F., Sower, L.L., Mahany, P.G., and Drummond, P.C. 1974. (Z,E)-7,11-hexadecadien-1-ol acetate: the sex pheromone of the angoumois grain moth, Sitotroga cerealella. Experientia. 30:17-18.
Voerman, S., and Rothschild, G.H.L. 1978c. Synthesis of the two components of the sex pheromone system of the potato tuberworm moth, Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae) and field experience with them. J. Chem. Ecol. 4:531-542.
  Walgenbach, C.A., Phillips, J.K., Burkholder, W.E., King, G.G.S., Slessor, K.N., and Mori, K. 1987. Determination of chirality in 5-hydroxy-4-methyl-3-heptanone, the aggregation pheromone of Sitophilus oryae (L.) and S. zeamais Motschulsky. J. Chem. Ecol. 13:2159-2169.
White, P.R., Chambers, J., Walter, C.M., Wilkins, J.P.G., and Millar, J.G. 1989. Saw-toothed grain beetle Oryzaephilus surinamensis (L.) (Coleoptera: Silvanidae). Collection, identification, and bioassay of attractive volatiles from beetles and oats. J. Chem. Ecol. 15: 999-1013.
 Williams, H.J., Silverstein, R.M., Burkholder, W.E., and Khorramshahi, A. 1981a. Dominicalure 1 and 2: components of aggregation pheromone from male lesser grain borer Rhyzopertha dominica (F.) (Coleoptera: Bostrichidae). J. Chem. Ecol. 7:759-784.

Yamaoka, R., Fukami, H., and Ishii, S. 1976. Isolation and identification of the female sex pheromone of the potato tuberworm moth, Phthorimaea operculella (Zeller). Agric. Biol. Chem. 40:1971-1977.