Photoperiod on biology

Natural day-night changes in light intensity and seasonal changes in day length influence stored-product insect physiology, behavior, and ecology (see Table 22.4 in Fundamentals of Stored Product Entomology). Table gives examples of changes in light intensity either directly initiating periods of feeding, development, diapause, locomotion, pheromone release, mating or oviposition, or entraining an internal endogenous biorhythm for these activities. The following list adds 114 studies to the 35 studies listed in Table 22.4. Gottlieb (2019) discussed the use of chronobiology in the management of stored product insect pests.

Effects of photoperiod on development (48), pheromone and mating (26), diapause (18), oviposition (16), locomotion (12), flight (8), susceptibility ionizing radiation or insecticide (7), reproductive diapause (5), oxygen consumption (5), circadian clock (4), sperm production and release (4) and parasitization (2) were the subject of the number of studies indicated in parentheses. Effects of photoperiod on death-feigning, egg cannibalism, exposure-concealment, feeding, geotaxis, hatching, larval penetration, longevity, productivity and survival, were each considered by one study. Several studies show that insects can respond to thermoperiod like they do to photoperiod.

A total of 56 species have been studied, most by fewer than 6 studies, but Plodia interpunctella (24 studies), Anthrenus verbasci (16 studies), Ephestia kuhniella (16 studies) have been studied more.

Studies on photoperiod or thermoperiod have increased over time: 2 from 1940-1949, 2 from 1950-1959, 8 from 1960-1969, 40 from 1970-1979, 19 from 1980-1989, 24 from 1990-1999, 29 from 2000-2009 and 25 from 2010-2019.


Abdel-Kader, M., Abdu, R.M., Husseion, M.A., 1987. Effect of time of day and temperature on sex pheromone production and perception by the rust-red flour beetle. Arab Gulf J. Sci. Res. Agric. Biol. Sci. B5: 147-156. (pheromone, mating)

Allotey, J., Morris, J.G., 2013. Feeding and mating activity patterns of stored product beetle, Cathartus quadricollis (Guerin-Meneville) (Coleoptera: Silvanidae). J. Appl. Zool. Res. 24: 109-112.

Alrubeai, H. F. 1987. The influence of temperature and photoperiod on development and reproduction of Ephestia calidella. Date Palm J. 5: 188-198. (development, oviposition)

Arai, T. 1979. Effects of light-dark cycles on the diel rhythm of several behaviours in the Indian meal moth, Plodia interpunctella Hubner (Lepidoptera: Pyralidae). Jap. J. Ecol. 29: 273-279. (mating, oviposition, development, locomotion)

Aslam, M., D. Hagstrum and B. Dover. 1994. The Effect of Photoperiod on the Flight Activity and Biology of Rhyzopertha dominica (Coleoptera, Bostrichidae). J. Kans. Entomol. Soc. 67:107-115. (flight)

Bakker, K. et al. 1963. On the relations between the duration of the larval and pupal period, weight and diurnal rhythm in emergence in Drosophila melanogaster. Entomol. Exp. Appl. 6: 37-52. (development)

Bashir, T., L. A. Birkinshaw, D. Farman, D. R. Hall and R. J. Hodges. 2003. 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. Journal of Stored Products Research 39(2): 159-169. (pheromone, mating)

Bell, C. 1977. Sensitivity of Larval Plodia interpunctella and Ephestia elutella (Lepidoptera) to Light during Photoperiodic Induction of Diapause. Physiol. Entomol. 2:167-172. (diapause)

Bell, C. 1983. The Regulation of Development during Diapause in Ephestia elutella (Hubner) by Temperature and Photoperiod. J. Insect Physiol. 29:485-490. (development)

Bell, C. H. 1981. The Influence of Light Cycle and Circadian-Rhythm on Oviposition in 5 Pyralid Moth Pests of Stored Products. Physiol. Entomol. 6:231-239. (oviposition)

Bell, C., and C. Bowley. 1980. Effect of Photoperiod and Temperature on Diapause in a Florida Strain of the Tropical Warehouse Moth Ephestia cautella. J. Insect Physiol. 26:533-538. (diapause)

Bell, C.H. 1991. Activity rhythms linked with foraging behaviour in insecticide-resistant and susceptible strains of Oryzaephilus surinamensis (Coleoptera: Silvanidae). Journal of Stored Products Research 27: 171–177. (locomotion)

Bell, C.H. and Kerslake, P.R. 1986. A circadian rhythm influencing foraging behaviour in the saw-toothed grain beetle Oryzaephilus surinamensis. Physiological Entomology 11: 1–6. (locomotion)

Bell, C.H. and Walker, D.J. 1973. Diapause induction in Ephestia elutella (Hübner) and Plodia interpunctella (Hübner) (Lepidoptera, Pyralidae) with a dawn-dusk lighting system. J. Stored Prod. Res. 9(3): 149-158. (diapause)

Bentley, E. W., D. L. Gunn, and D. W. Ewer. 1941. The Biology and Behaviour of Ptinus tectus Boie. (Coleoptera, Ptinidae), a Pest of Stored Products. I. The daily rhythm of locomotory activity, especially in relation to light and temperature. J. Exp. Biol. 18:182-195. (locomotion)

Burks, C. S., D. G. Brandl, and B. S. Higbee. 2011. Effect of natural and artificial photoperiods and fluctuating temperature on age of first mating and mating frequency in the navel orangeworm, Amyelois transitella. J. Insect Sci. 11:48-48. (pheromone, mating)

Campbell, B.O., 1964. Solar and Lunar Periodicities in Oxygen Consumption by the Mealworm, Tenebrio molitor. Ph.D. thesis. Northwestren University (oxygen consumption)

Chen, H., H. Zhang, K. Y. Zhu, and J. E. Throne. 2012. Induction of reproductive diapause in Habrobracon hebetor (Hymenoptera: Braconidae) when reared at different photoperiods at low temperatures. Environ. Entomol. 41:697-705. (reproductive diapause)

Chiba, Y., L. K. Cutkomp, and F. Halberg. 1974. Circadian oxygen consumption rhythm of the flour beetle, Tribolium confusum with special reference to phase-shifting. p. 602-606. In Chronobiology. (oxygen consumption)

Chiba, Yoshihiko, L. K. Cutkomp and Franz Halberg. 1973. Circadian oxygen consumption rhythm of the flour beetle, Tribolium confusum. Journal of insect physiology 19(11): 2163-2172. (oxygen consumption)

Cloudsley‐Thompson, J. L. 1953. Studies in diurnal rhythms. iv. photoperiodism and geotaxis in Tenebrio molitor L. (Coleoptera: Tenebrionidae). In Proceedings of the Royal Entomological Society of London. Series A, General Entomology 28(10‐12): 117-132.

Coffelt, J.A., Sower, L.L. and Vick, K.W. 1978. Quantitative analysis of identified compounds in pheromone gland rinses of Plodia interpunctella and Ephestia cautella at Different Times of Day. Environ. Entomol. 7 (4), 502-505. (pheromone, mating)

Cox, P. 1975. Influence of Photoperiod on Life-Cycles of Ephestia calidella (Guenee) and Ephestia figulilella Gregson (Lepidoptera-Phycitidae). J. Stored Prod. Res. 11:75-85. (diapause)

Cox, P. D. 1979. The influence of photoperiod on the life-cycle of Ectomyelois ceratoniae (Zeller) (Lepidoptera: Pyralidae). J. Stored Prod. Res. 15:111-115.

Cymborowski, B. 1977. Effect of Lighting Conditions on Growth and Developmental Rhythms of Singly Reared Dermestes vulpinus (F). Comparative Biochemistry and Physiology A-Physiology 58:99-101. (Dermestes maculatus, development)

Cymborowski, B., and J. Giebultowicz. 1976. Effect of Photoperiod on Development and Fecundity in Flour Moth, Ephestia kuehniella. J. Insect Physiol. 22:1213-1217.

Edwards, D. K. 1962. Laboratory determinations of the daily flight times of separate sexes of some moths in naturally changing light. Can. J. Zool. 40:511-530. (flight)

Ewer, R. F. 1943. Diurnal activity of three insect pests of stored products. Nature 152:133-134. (locomotion)

Fexova, S., 2010. Circadian Clock of Two Insect Model Species – Drosophila melanogaster and Tribolium castaneum. Master Thesis. University of South Bohemia Faculty of Science

Fleming, D. A. 2008. The influence of photoperiod upon the productivity of Oryzaephilus surinamensis (L.) (Coleoptera: Silvanidae). J. Stored Prod. Res. 44:213-218.

Fondacaro, J. D., and A. Butz. 1970. Circadian Rhythm of Locomotor Activity and Susceptibility to Methyl Parathion of Adult Tenebrio molitor (Coleoptera – Tenebrionidae). Ann. Entomol. Soc. Am. 63:952-955. (locomotion, susceptibility)

Giebultowicz, J., and B. Cymborowski. 1976. Effect of Different Lighting Conditions on Circadian Developmental Rhythms and Behavior in Ephestia kuehniella Zell. Journal of Interdisciplinary Cycle Research 7:119-126. (development)

Gottlieb, Daphna 2019. Agro-chronobiology: Integrating circadian clocks /time biology into storage management. J. Stored Prod. Res. 82: 9-16.

Guerber, J. C. 1971. Photoperiodic mating rhythm for the angoumois grain moth, Sitotroga cerealella (Lepidoptera: Gelechiidae), and the role of male response to the female sex pheromone in its determination. MS Thesis, Kansas State University Department of Entomology (pheromone, mating)

Hagstrum, D. W. and Tomblin, C. F.  1973.  Oviposition by the almond moth Cadra cautella in response to falling temperature and onset of darkness.  Ann. Entomol. Soc. Am. 66: 809‑812. (oviposition)

Hammack, L. 1974. Sex pheromone-releasing behavior of Trogoderma glabrum females in relation to sex pheromone production, glandular location, mating behavior, and photoperiod; with comparative notes on six other Trogoderma species (Coleoptera: Dermestidae). Ph.D. dissertation, University of Wisconsin, Madison (pheromone, mating)

Hammack, L., and Burkholder, W. E. 1981. Calling behaviour in female Trogoderma granarium Everts (Coleoptera: Dermestidae). J. Stored Prod. Res. 17:25-29. (pheromone, mating)

Hammack, L., and W. Burkholder. 1976. Circadian-Rhythm of Sex Pheromone-Releasing Behavior in Females of Dermestid Beetle, Trogoderma glabrum – Regulation by Photoperiod. J. Insect Physiol. 22:385-388. (pheromone, mating)

Hammack, L., M. Ma, and W. Burkholder. 1976. Sex Pheromone-Releasing Behavior in Females of Dermestid Beetle, Trogoderma glabrum. J. Insect Physiol. 22:555-561. (pheromone, mating)

Harano, T., and T. Miyatake. 2010. Genetic basis of incidence and period length of circadian rhythm for locomotor activity in populations of a seed beetle. Heredity 10:268-273. (locomotion)

Harano, T., and T. Miyatake. 2011. Independence of genetic variation between circadian rhythm and development time in the seed beetle, Callosobruchus chinensis. J. Insect Physiol. 57:415-420. (development)

Hemmati, C., S. Moharramipour, and A. A. Talebi. 2017. Diapause Induced by Temperature and Photoperiod Affects Fatty Acid Compositions and Cold Tolerance of Phthorimaea Operculella (Lepidoptera: Gelechiidae). Environ. Entomol. 46:1456-1463.

Higashi, T., M. Takeda, and S. Momoi. 2005. Temperature and photoperiodic regulation of the postembryonic development of the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae). Japanese Journal of Environmental Entomology and Zoology 16:1-7. (development)

Hossain, T., M. Yasmin, M. H. Islam, Islam, A. T. M. F., and A. S. M. Saifullah. 2016. Effects of photoperiod on the development of hide beetle, Dermestes maculatus DeGeer (Coleoptera: Dermestidae). Journal of Entomology and Zoology Studies 4:672-676. (development)

Ismail, I. I., El-Nahal, A. K. M., Kamel, A. H., and Mostafa, T. S. 1988. Effect of light on the development of the Angoumois grain moth, Sitotroga cerealella (Olivier) (Lepidoptera: Gelechiidae). Insect Sci. Appl. 9:27-29. (development)

Kikukawa, S., A. Higuchi, T. Ikeda, T. Mukai, H. Okamoto, K. Tanaka, and S. Yaguchi. 2009. Determination of effective light pulse and classical Bunsow experiment in the larval diapause of the Indian meal moth Plodia interpunctella. Physiol. Entomol. 34:333-337. (diapause)

Kikukawa, S., and K. Ohde. 2007. The role of the main photophase on dark-time measurement used for diapause determination in the Indian meal moth, Plodia interpunctella. Physiol. Entomol. 32:351-356. (diapause)

Kikukawa, S., and S. Masaki. 1984. Interacting Effects of Photophase and Scotophase on the Diapause Response of the Indian Meal Moth, Plodia-Interpunctella. J. Insect Physiol. 30:919-925. (diapause)

Kikukawa, S., H. Kubota, H. Ohkouchi, and K. Tateiwa. 1998. The effect of temperature and light pulses on the induction of diapause in the Toyama strain of the Indian meal moth, Plodia interpunctella. Physiol. Entomol. 23:249-254. (diapause)

Kikukawa, S., H. Nakamura, A. Saitoh, and K. Terayama. 2016. Effect of thermoperiod amplitude on the adult eclosion rhythm of Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae). Entomol. Sci. 19:310-311. (development)

Kikukawa, S., R. Hashizume, M. Honda, Y. Inoue, T. Maekawa, M. Miyabayashi, and N. Mori. 2012. Effects of photoperiod and temperature on the rhythm and free‐running of adult eclosion in the Indian meal moth Plodia interpunctella. Physiol. Entomol. 37:258-265. (development)

Kikukawa, S., R. Hashizume, M. Honda, Y. Inoue, T. Maekawa, R. Sakata, N. Takahashi, K. Tanaka, and Y. Uchida. 2013. Adult eclosion rhythm of the Indian meal moth Plodia interpunctella: Response to various thermocycles with different means and amplitudes. Physiol. Entomol. 38:253-259. (development)

Kikukawa, S., R. Rou, and M. Sugimoto. 2005. Effect of light pulses in early scotophase on resetting of the night-measuring diapause clock of the Indian meal moth, Plodia interpunctella. Physiol. Entomol. 30:256-261. (diapause)

Kikukawa, S., T. Minamizuka, and W. Matoba. 2008. Responses to stepwise photoperiodic changes for the larval diapause of the Indian meal moth Plodia interpunctella. Physiol. Entomol. 33:360-364. (diapause)

Kikukawa, S., Y. Arakawa, K. Hayakawa, M. Hayashi, K. Katou, J. Kaneshige, M. Kimura, T. Nakamura, Y. Nakamura, and H. Watanabe. 2009. Effects of skeleton photoperiods on the induction of larval diapause in the Indian meal moth Plodia interpunctella. Physiol. Entomol. 34:180-184. (diapause)

Kikukawa, S., Y. Kakihara, Y. Okano, R. Shindou, N. Sugino, J. Tsunekawa, A. Yasui, and K. Yoneda. 2015. Eclosion Rhythm of Plodia interpunctella Under Non-24 h Thermocycles. International Journal of Animal Biology 1:273-280. (development)

Kobelkova, A., Zavodska, R., Sauman, I., Bazalova, O., Dolezel, D., 2015. Expression of clock genes period and timeless in the central nervous system of the Mediterranean Flour Moth, Ephestia kuehniella. J. Biol. Rhythm. 30(2): 104-116.

Kobelková, Alena, Radka Závodská, Ivo Sauman, Olga Bazalová, and David Dolezel. 2015. Expression of clock genes period and timeless in the central nervous system of the Mediterranean flour moth, Ephestia kuehniella. Journal of biological rhythms 30(2): 104-116.

Kurota, H., and M. Shimada. 2003. Photoperiod-dependent adult diapause within a geographical cline in the multivoltine bruchid Bruchidius dorsalis. Entomol. Exp. Appl. 106:177-185. (reproductive diapause)

Landolt, P. J., and C. E. Curtis. 1982. Effects of temperature on the circadian rhythm of navel orange worm sexual activity. Environ. Entomol. 11:107-110. (pheromone, mating)

Lenga, A., and J. Huignard. 1992. Effect of changes in the thermoperiod on reproductive diapause in Bruchidius atrolineatus Pic (Coleoptera: Bruchidae). Physiol. Entomol. 17:247-254. (reproductive diapause)

Lenga, A., C. Thibeaudeau, and J. Huignard. 1991. Influence of thermoperiod and photoperiod on reproductive diapause in Bruchidius atrolineatus (Pic) (Coleoptera, Bruchidae). Physiol. Entomol. 16:295-303. (reproductive diapause)

Lenga, A., I. Glitho, and J. Huignard. 1993. Interactions between photoperiod, relative humidity and host-plant cues on the reproductive diapause tennination in Bruchidius atrolineatus Pic (Coleoptera, Bruchidae). Invertebrate reproduction and development 24:87-96. (reproductive diapause)

Li, Cheng‐Jun, Xiao‐Pei Yun, Xiao‐Juan Yu and Bin Li. 2018. Functional analysis of the circadian clock gene timeless in Tribolium castaneum. Insect science 25(3): 418-428.

Limonta, L., M. Stampini, and D. P. Locatelli. 2010. Development of rusty wave (Idaea inquinata) at constant temperatures, relative humidities and photoperiods. Bulletin of Insectology 63:171-174. (development)

Lovitt, A. E., and Soderstrom, E. L. 1973. Effect of varying light and temperature cycles on the ovipositional response of malathion-treated Indian meal moth adults. J. Econ. Entomol. 66:167-170. (oviposition)

Lum, P. T. M., and Flaherty, B. R. 1970. Regulating oviposition by Plodia interpunctella in the laboratory by light and dark conditions. J. Econ. Entomol. 63:236-239. (oviposition)

Lum, P. T. M., and Flaherty, B. R. 1973. Influence of continuous light on oöcycte maturation in Bracon hebetor. Ann. Entomol. Soc. Am. 66:355-357.

Madrid, F. J., and Sinha, R. N. 1983. Movement and oviposition of Ephestia cautella (Walker) and Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae) of different ages in response to seasonal light changes. Can. J. Zool. 61:1726-1732.

Matsuno, T., Y. Kawasaki, and H. Numata. 2013. Small geographic variation in photoperiodic entrainment of the circannual rhythm in the varied carpet beetle, Anthrenus verbasci. Zool. Sci. 30:304-310.

Matsuno, T., Y. Miyazaki, N. Muramatsu, and H. Numata. 2013. Circannual pupation timing is not correlated with circadian period in the varied carpet beetle Anthrenus verbasci. Biol. Rhythm Res. 44:849-855. (development)

Mbata, G. N. 1985. Some physical and biological factors affecting oviposition by Plodia interpunctella (Hübner) (Lepidoptera: Phycitidae). Insect Sci. Appl. 6:597-604. (oviposition)

Mbata, G., S. Shu, and S. Ramaswamy. 1997. Rhythmicity of mating and oviposition in Callosobruchus subinnotatus (Pic) (Coleoptera: Bruchidae). J. Insect Behav. 10:409-423.

Melou, J.F. 1985. Variability of sensibility to parathion in relation to the exposure hour of a wild strain of Drosophilae melanogaster from Benin. Acta Oecol (Oecol Appl), 6:15-21. (susceptibility ionizing radiation or insecticide)

Miyatake, T. 2001. Diurnal periodicity of death-feigning in Cylas formicarius (Coleoptera: Brentidae). J. Insect Behav. 14:421-432.

Miyazaki, Y., and H. Numata. 2009. Responsiveness to photoperiodic changes in the circannual rhythm of the varied carpet beetle, Anthrenus verbasci. Journal of Comparative Physiology A-Neuroethology Sensory Neural and Behavioral Physiology 195:241-246. (development)

Miyazaki, Y., and H. Numata. 2010. Exhibition of circannual rhythm under constant light in the varied carpet beetle Anthrenus verbasci. Biol. Rhythm Res. 41:441-448. (development)

Miyazaki, Y., H. Numata, and T. Nisimura. 2004. Phase response to long-day pulses under natural photoperiod in the circannual rhythm of Anthrenus verbasci. Zool. Sci. 21:1324-1324. (development)

Miyazaki, Y., H. Numata, and T. Nisimura. 2005. Involvenment of the circadian system in the photoperiodic time measurement for entrainment of the circannual rhythm in Anthrenus verbasci. Zool. Sci. 22:1491-1491. (development)

Miyazaki, Y., H. Numata, and T. Nisimura. 2006. Long-day stimulus that induces arrhythmicity in the circannual pupation rhythm of Anthrenus verbasci. Zool. Sci. 23:1194-1194. (development)

Miyazaki, Y., T. Nisimura, and H. Numata. 2005. A phase response curve for circannual rhythm in the varied carpet beetle Anthrenus verbasci. Journal of Comparative Physiology A-Neuroethology Sensory Neural and Behavioral Physiology 191:883-887. (development)

Miyazaki, Y., T. Nisimura, and H. Numata. 2006. Phase responses in the circannual rhythm of the varied carpet beetle, Anthrenus verbasci, under naturally changing day length. Zool. Sci. 23:1031-1037. (development)

Miyazaki, Y., T. Nisimura, and H. Numata. 2009. A circadian system is involved in photoperiodic entrainment of the circannual rhythm of Anthrenus verbasci. J. Insect Physiol. 55:494-498. (development)

Miyazaki, Y., T. Nisimura, and H. Numata. 2009. Circannual pupation rhythm in the varied carpet beetle Anthrenus verbasci under different nutrient conditions. Entomol. Sci. 12:370-375. (development)

Miyazaki, Yosuke, Tomoyosi Nisimura, and Hideharu Numata. 2007. Phase resetting and phase singularity of an insect circannual oscillator. Journal of Comparative Physiology A 193(11): 1169-1176. (Anthrenus verbasci, development)

Miyazaki, Yosuke, Tomoyosi Nisimura, and Hideharu Numata. 2012. Circannual rhythm in the varied carpet beetle, Anthrenus verbasci. Progress in brain research 199: 439-456. (development)

Moriarty, F. 1959. The 24-Hr Rhythm of Emergence of Ephestia kuhniella Zell from the Pupa. J. Insect Physiol. 3:357-366. (development)

Ndoutoume-Ndong, A., D. Rojas-Rousse, and R. Allemand. 2006. Rythmes d’activité locomotrice chez deux insectes parasitoïdes sympatriques: Eupelmus orientalis et Eupelmus vuilleti (Hyménoptère, Eupelmidae) (Locomotor activity rhythms in two sympatric parasitoid insects: Eupelmus orientalis and Eupelmus vuilleti (Hymenoptera, Eupelmidae)). Comptes Rendus Biologies 329:476-482. (locomotion)

Neven, L. G. 2013. Effects of Short Photoperiod on Codling Moth Diapause and Survival. Environ. Entomol. 42:520-523.

Nielson, M. W. 1976. Diapause in the alfalfa seed chalcid, Bruchophagus roddi (Gussakovsky) in relation to natural photoperiod. Environ. Entomol. 5:123-127. (diapause)

Nisimura, T., and H. Numata. 2001. Endogenous timing mechanism controlling the circannual pupation rhythm of the varied carpet beetle Anthrenus verbasci. Journal of Comparative Physiology A-Sensory Neural and Behavioral Physiology 187:433-440. (development)

Nisimura, T., and H. Numata. 2002. Evaluation of the frequency demultiplication hypothesis of circannual pupation rhythm in the varied carpet beetle Anthrenus verbasci (Coleoptera : Dermestidae). Biol. Rhythm Res. 33:255-260. (development)

Noble Singh, V. T., S. Sam Manohar Das. 2016. Chronobiological Management of Pulse Beetle Callosobruchus maculatus Fab. (Coleoptera: Bruchidae). International Journal of Advances in Science Engineering and Technology 4(3): 117-121. (susceptibility ionizing radiation or insecticide)

Nordlund, D. A., and Brady, U. E. 1974a. The calling behavior of female Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae) under two light regimes. Environ. Entomol. 3:793-796. (pheromone, mating)

Nordlund, D. A., and Brady, U. E. 1974b. Factors affecting release rate and production of sex pheromone by female Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae). Environ. Entomol. 3:797-798. (pheromone, mating)

Nowock, J. 1973. Effect of Farnesyl Methyl-Ether on Molting Rhythm and Testes Development in Ephestia kuhniella – Studies In-Vivo and In-Vitro. Wilhelm Roux Archiv Fur Entwicklungsmechanik Der Organismen 172:303-316.

Numata, H., Y. Miyazaki, and T. Nisimura. 2004. Entrainment of circannual rhythms: An example in the varied carpet beetle Anthrenus verbasci. Zool. Sci. 21:1238-1238. (development)

Obeng-Ofori, D., and T. H. Coaker. 1990. Some factors affecting responses of four stored product beetles (Coleoptera: Tenebrionidae & Bostrichidae) to pheromones. Bulletin of Entomological Research 80(4): 433-441. (pheromone, mating)

O’ceallachain, D. P., and M. F. Ryan. 1977. Production and perception of pheromones by the beetle Tribolium confusum. Journal of Insect Physiology 23(10): 1303-1309. (pheromone, mating)

Odeyemi, O., and A. Hassan. 1993. Influence of Temperature, Humidity and Photoperiod on Oviposition and Larval Development in Trogoderma granarium Everts (Coleoptera, Dermestidae). Appl. Entomol. Zool. 28:275-281.  (oviposition)

Okiwelu, S., P. Dudu and N. Lale. 1998. The effects of photoperiod on some aspects of the biology of Oryzaephilus mercator (Fauvel) (Coleoptera : Silvanidae). J. Stored Prod. Res. 34:189-193.

Ono,T.,Charlton,R.E.,and Carde,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

Pesson, P., and G. Girish. 1968. Sensitivity of Sitophilus zeamais Mots (=S oryzae L) to Ionizing Radiations at Different Stages of its Development. Radiographical Study of Endogenous Stages and Actographical Registration. Annales Des Epiphyties 19:513. (susceptibility ionizing radiation or insecticide)

Pizzol, J., and B. Pintureau. 2008. Effect of photoperiod experienced by parents on diapause induction in Trichogramma cacoeciae. Entomol. Exp. Appl. 127:72-77. (diapause)

Pristavko, V. P. et al. 1974. Circadian rhythm of the radiosensitivity of the codling moth Laspeyresia pomonella L. (Lepidoptera, Tortricidae). Radiobiology Radiobiologiia 13:159-162. (susceptibility ionizing radiation or insecticide)

Quartey, G., and T. Coaker. 1996. Periodicity of activity and sexual response of adult Ephestia cautella (Walk) (Lep, Phycitidae). Journal of Applied Entomology-Zeitschrift Fur Angewandte Entomologie 120: 335-339. (flight)

Rafaeli, A., Gileadi, C., 1995. Factors affecting pheromone production in the stored product moth, Plodia interpunctella a preliminary study. J. Stored Prod. Res. 31(3): 243-247 (pheromone, mating)

Rakowski, G., and B. Cymborowski. 1982. Aggregation Pheromone in Dermestes maculatus – Effects on Larval Growth and Developmental Rhythms. International Journal of Invertebrate Reproduction 4:249-254. (development)

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