Resource-locating behavior

Mobility allows insects to locate essential resources (food, shelter, mate, oviposition site and favorable environment) and avoid natural enemies or unfavorable environments. Searching generally involves multi-sensory orientation (odor, tactile, visual, etc) which may be modified by insect physiological state (age, sex, mating or feeding history, hydration). Insects often choose the most favorable location in a chemical, dockage, humidity or temperature gradient. The following references expand the discussion on pages 51-53 of Fundamentals of Stored Product Entomology. Some of the sensilla detecting behavioral cues have been studied for 34 species of stored-product insects.


Barton Browne, L. 1993. Physiologically induced changes in resource-oriented behavior. Annual Rev. Entomol. 38:1-25.

Bell, C. H. 2014. A review of insect responses to variations encountered in the managed storage environment. Journal of Stored Products Research 59: 260-274

Campbell, J. F. 2006. Stored-product insect behavior, pp. 39-51. In J. W. Heaps (ed.), Insect Management for Food Storage and Processing, 2nd ed. AACC International, St Paul, MN.

Cox, P. and L. Collins. 2002. Factors affecting the behaviour of beetle pests in stored grain, with particular reference to the development of lures. J. Stored Prod. Res. 38:95-115.

Gadenne, Christophe, Romina B. Barrozo and Sylvia Anton. 2016. Plasticity in Insect Olfaction: To Smell or Not to Smell? Annu. Rev. Entomol. 61: 317–333.

Longstaff, B. 1981. Biology of the Grain Pest Species of the Genus Sitophilus (Coleoptera, Curculionidae) – a Critical-Review. Protection Ecology 2: 83-130.

Mult-sensory orientation

Amos, T. G. 1968. The effect of light on the humidity reactions of Carpophilus dimidiatus and Tribolium castaneum. Entomologia Experimentalis et Applicata 11(3): 331-340

Arbogast, R. T. and Carthon, M. 1970. Light, tactile and humidity responses of larvae of Oryzaephilus surinamensis (Coleoptera: Cucujidae). Entomol. Exp. Appl. 13: 395-402.

Arbogast, R. T. and B. R. Flaherty. 1973. Light, tactile, and humidity responses of adult Oryzaephilus surinamensis. Environ. Entomol. 2: 931-935.

Armitage, D. M., Day, P. G., and Lewis, G. J. 1983. The effect of aeration on the vertical distribution of grain beetles (Coleoptera) in wheat. Bull. Entomol. Res. 73: 233-237.(humidity, vibration and wind speed)

Arnold, Sarah E. J., Philip, C. Stevenson and Steven R. Belmain. 2016. Shades of yellow: interactive effects of visual and odour cues in a pest beetle. PeerJ 4: e2219. (Sitophilus zeamais)

Arnold, Sarah E. J., Stevenson, Philip C., Belmain, Steven R., 2015. Responses to colour and host odour cues in three cereal pest species, in the context of ecology and control. Bull. Entomological Res. 105(4): 417-425. (Rhyzopertha dominica, Prostephanus truncatus, Sitophilus zeamais)

Bartels, D. 2000. Dispersion behaviour of Oryzaephilus mercator (Fauvel) (Coleoptera: Silvanidae) in phototaxis, geotaxis and thigmotaxis. IOBC/wprs Bulletin 23: 227-231.

Browning, H. C. 1946, 1947. Mechanical disturbance and light as factors influencing the pullulation of Calandra granaria Linn. Proceedings of the Zoological Society of London 116(3-4): 675-691.

Campbell, J. F. 2012. Attraction of walking Tribolium castaneum adults to traps. Journal of Stored Products Research 51: 11-22. (food odor, pheromone and wind)

Cox, P. D. and Parish, W. E. 1991. Effects of refuge content and food availability on refuge-seeking behavior in Cryptolestes ferrugineus (Stephens) (Coleoptera: Cucujidae), J. Stored Prod. Res. 27: 135-139.

Dowdy, A. K. 1994. Flight initiation of lesser grain borer (Coleoptera: Bostrichidae) as influenced by temperature, humidity and light. J. Econ. Entomol. 87: 1714-1717.

Duehl, Adrian J., Cohnstaedt, L.W., Arbogast, R.T., Teal, P.E.A. 2011. Evaluating light attraction to increase trap efficiency for Tribolium castaneum (Coleoptera: Tenebrionidae). Journal of Economic Entomology 104: 1430-1435. (pheromone)

Fadamiro, H. Y. and Wyatt, T. D. 1995. Flight initiation by Prostephanus truncatus in relation to time of day, temperature, relative humidity and starvation. Entomol. Exp. Appl. 75: 273-277.

Fadamiro, H., T. Wyatt and M. Birch. 1998. Flying beetles respond as moths predict: Optomotor anemotaxis to pheromone plumes at different heights. J. Insect Behav. 11:549-557.

Fouad, Hany Ahmed, Lêda Rita D’Antonino Faroni, Evaldo Ferreira Vilela, and Eraldo Rodrigues de Lima. 2013. Flight responses of Sitotroga cerealella (Lepidoptera: Gelechiidae) to corn kernel volatiles in a wind tunnel. Arthropod-Plant Interactions 7(6): 651-658. (wind, food odor)

Karlis, G., C. G. Athanassiou and C. Th Buchelos. 2008. Monitoring insect populations by using adhesive surfaces of different colours in a dried fig warehouse in Southern Greece. IOBC/wprs Bulletin 40:  93-106. (pheromone and light, Plodia interpunctella, Habrobracon hebetor)

Kennedy, J. S. and D. Marsh. 1974. Pheromone-regulated anemotaxis in flying moths. Science 184: 999-1002.

Kuenen, L. P. S., and H. C. Rowe. 2006. Cowpea weevil flights to a point source of female sex pheromone: analyses of flight tracks at three wind speeds. Physiological Entomology 31(2): 103-109.

Levinson, H. and T. Hoppe. 1983. Preferential Flight of Plodia-Interpunctella and Cadra-Cautella (Phycitinae) Toward Figures of Definite Shape and Position with Notes on the Interaction between Optical and Pheromone Stimuli. Zeitschrift Fur Angewandte Entomologie-Journal of Applied Entomology 96: 491-500.

Mankin, R. W., K. W. Vick, M. S. Mayer and J. A. Coffelt. 1980. Anemotactic response threshold of the Indian meal moth, Plodia interpunctella (Hübner)(Lepidoptera: Pyralidae), to its sex pheromone.” Journal of Chemical Ecology 6(5): 919-928.

Marsh, D., J. S. Kennedy and A. R. Ludlow. 1978. An analysis of anemotactic zigzagging flight in male moths stimulated by pheromone. Physiol. Entomol. 3: 221-240. (Plodia interpunctella)

Mathieu, F., Brun, L . O., Marcillaud, C. and Frérot, B. 1997. Trapping of the coffee berry borer Hypothenemus hampei Ferr. (Col., Scolytidae) within a mesh-enclosed environment: interaction of olfactory and visual stimuli. J. Appl. Entomol. 121: 181-186.

McQuate, Grant T. 2014. Green light synergistally enhances male sweetpotato weevil response to sex pheromone. Scientific reports 4: 4499.

Miyatake, Takahisa, Tomoyuki Yokoi, Taro Fuchikawa, Nobuyoshi Korehisa, Toru Kamura, Kana Nanba, Shinsuke Ryouji et al. 2016. Monitoring and Detecting the Cigarette Beetle (Coleoptera: Anobiidae) Using Ultraviolet (LED) Direct and Reflected Lights and/or Pheromone Traps in a Laboratory and a Storehouse. Journal of Economic Entomology 109(6): 2551-2560.

Nakamoto, H. and J. Takushi. 2001. Positive phtotactic response of the West Indian sweet potato weevil, Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) to chemiluminescent light and effectiveness of the light source for attracting weevils in the field. Jpn. J. Appl. Entomol. Zool. 45: 212–214. (food odor and light)

Nansen, Christian, Thomas W. Phillips, and Stacy Sanders. 2004. Effects of height and adjacent surfaces on captures of Indianmeal moth (Lepidoptera: Pyralidae) in pheromone-baited traps. Journal of economic entomology 97(4): 1284-1290.

Nawrot, Jan, Magdalena Gawlak, Janusz Szafranek, Beata Szafranek, Elżbieta Synak, Jerzy R. Warchalewski, Dorota Piasecka-Kwiatkowska, Wioletta Błaszczak, Tomasz Jeliński and Józef Fornal. 2010. The effect of wheat grain composition, cuticular lipids and kernel surface microstructure on feeding, egg-laying, and the development of the granary weevil, Sitophilus granarius (L.). Journal of stored products research 46(2): 133-141.

Obeng-Ofori, D. 1991. Analysis of Orientation Behavior of Tribolium castaneum and T. confusum to Synthetic Aggregation Pheromone. Entomol. Exp. Appl. 60: 125-133. (pheromone and wind)

Parde, S. R., Jayas, D. S. and White, N. D. G. 2004. Movement of Cryptolestes ferrugineus (Coleoptera: Cucujidae) in grain columns containing pockets of high moisture content wheat and carbon dioxide gradients. J. Stored Prod. Res. 40: 299-316.

Parra, J., S. Vinson, S. Gomes and F. Consoli. 1996. Flight response of Habrobracon hebetor (Say) (Hymenoptera: Braconidae) in a wind tunnel to volatiles associated with infestations of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae). Biological Control 6: 143-150.

Perttunen, V. and M. Lahermaa. 1963. The light reactions of the larvae and adults of Tenebrio molitor L.(Col., Tenebrionidae) and their interference with the humidity reactions. Ann. Ent. Fenn. 29: 83-106.

Perttunen, V., and L. Paloheimo. 1963. Effect of temperature and light intensity on light reactions of larvae and adults of Tenebrio molitor. Ann. Entomol. Fenn. 29: 171-184.

Perttunen, V. 1973. Individual variation in the response of Rhyzopertha dominica F. (Col., Bostrichidae) to air humidity and light. Annales Entomologici Fennici 39: 129-142.

Phillips, Thomas W. and Michael R. Strand. 1994. Larval secretions and food odors affect orientation in female Plodia interpunctella. Entomologia Experimentalis et Applicata 71(3): 185-192.

Quartey, G. and T. Coaker. 1992. The Development of an Improved Model Trap for Monitoring Ephestia cautella. Entomol. Exp. Appl. 64: 293-301. (pheromone, visual and wind)

Ra̧kowski, Grzegorz. 1988. Effect of illumination intensity on the response of the hide beetle, Dermestes maculatus, to aggregation pheromone. Journal of insect physiology 34(12): 1101-1104.

Romero, S.A., Campbell, J.F., Nechols, J.R. and With, K.A. 2010. Movement behavior of red flour beetle: response to habitat cues and patch boundaries. Environmental Entomology 39: 919-929. (air flow, shelter, food deprivation)

Sambaraju, Kishan R., and Thomas W. Phillips. 2008. Effects of physical and chemical factors on oviposition by Plodia interpunctella (Lepidoptera: Pyralidae). Annals of the Entomological Society of America 101(5): 955-963. (odor, size-shape kernel)

Semeao, A. A., J. F. Campbell, R. J. Whitworth, and P. E. Sloderbeck. 2011. Response of Tribolium castaneum and Tribolium confusum adults to vertical black shapes and its potential to improve trap capture. J. Stored Prod. Res. 47:88-94. (pheromones and kairomones)

Singh, Karan and S. K. Majumder. 1984. Studies on mate seeking behaviour and anemotactic response of Tropical Warehouse Moth, Ephestia cautella (Walker) to (Z, E)‐9, 12‐tetradecadienyl acetate. Journal of Applied Entomology 98(1-5): 317-320.

Smereka, E.P. and Hodson, A.C. 1959. Some humidity and light reactions of the granary weevil, Sitophilus granarius. Canadian Entomologist 91: 784–797.

Som, Gurcharn Singh 1986. Influence of food on the phototactic response of the larvae of Trogoderma granarium Everts (Coleóptera: Dermestidae). Indian Journal of Ecology, 13(1): 179-182.

Stejskal, V. 1995. The influence of food and shelter on the efficacy of a commercial sticky trap in Tribolium castaneum (Coleoptera: Tenebrionidae). J. Stored Prod. Res. 31: 229-233.

Weston, P. A. and Hoffman, S. A. 1991. Humidity and tactile responses of Sitophilus zeamais (Coleoptera: Curculionidae). Environ. Entomol. 20: 1433-1437.

Weston, P. A., and Rattingourd, P. L. 1999. Ovipositional stimuli of Angoumois grain moth (Lepidoptera: Gelechiidae), a primary pest of stored grain. J. Entomol. Sci. 34: 445-451. (grain chemical, crevice size)

Willis, E. B. and Roth, L. M. 1950a. Humidity reaction of Tribolium castaneum (Herbst). J. Exp. Zool. 115: 561-587. (food odor and humidity)

Willis, E. B. and Roth L. M. 1950b. The attraction of Tribolium castaneum to flour. J. Econ. Entomol. 43: 927-932. (food odor and humidity)

Willis, E. B. and Roth, L. M. 1954. Reactions of flour beetles of the genus Tribolium to carbon dioxide. J. Exp. Zool. 127: 117-152. (wind)

Wisniewska, J. and R. T. Carde. 2012. Visual cues collimate the trajectories of almond moth Cadra cautella males flying in wind and still air within a wind-formed plume of pheromone. Physiol. Entomol. 37: 42-52.

Wisniewska, J. 2001. Influence of visual cues in the flight of male almond moths (Cadra cautella) within plumes of sex pheromone. PhD Dissertation, University of California.

Modification by Physiological State

Adeesan, C., G. W. Rahalkar, and A. J. Tamhankar. 1969. Effect of Age and Previous Mating on Response of Khapra Beetle Males to Female Sex Pheromone. Entomol. Exp. Appl. 12: 229-234.

Amos, T. and F. Waterhouse. 1967. Effect of Desiccation on Light Reactions of Carophilus dimidiatus and Tribolium castaneum. Entomol. Exp. Appl. 10: 329-336.

Arbogast, R. and M. Carthon. 1972. Effect of Starvation and Desiccation on Water-Balance and Humidity Response of Adult Oryzaephilus surinamensis (Coleoptera-Cucujidae). Entomol. Exp. Appl. 15: 488-498.

Arnold, S.E.J., Stevenson, P.C. & Belmain, S.R. 2012. Odour-mediated orientation of beetles is influenced by age, sex and morph. PLoS ONE 7: e49071.

Aslam, M., Hagstrum, D. W. and Dover, B. A. 1994. The effect of photoperiod on the flight activity and biology of Rhyzopertha dominica (Coleoptera: Bostrichidae). J. Kans. Entomol. Soc. 67: 107-115.

Barlow, C. and Perttunen.V. 1966. Some Effects of Desiccation on Locomotion of Grain Beetle Sitophilus granarius (L) (Coleoptera-Curculionidae). Can. J. Zool. 44: 825-837.

Barrer, P. M., Starick, N. T., Morton, R. and Wright, E. J. 1993. Factors influencing initiation of flight by Rhyzopertha dominica (F.) (Coleoptera: Bostrichidae). J. Stored Prod. Res. 29: 1-5.

Bishara, Sadek Ibrahim. 1968. Comparison of some aspects of flight activity of Sitophilus zea-mais Mots. and Sitophilus oryzae L. Bull. Soc. Entomol. Egypte 52: 477-500.

Boughton, A. and H. Fadamiro. 1996. Effect of age and sex on the response of walking Prostephanus truncatus (Horn) (Coleoptera:Bostrichidae) to its male-produced aggregation pheromone. J. Stored Prod. Res. 32: 13-20.

Coffelt, J. A., and K. W. Vick. 1987. Sex pheromones of Ephestia cautella (Walker) (Lepidoptera Pyralidae): influence of mating on pheromone titer and release rate. Journal of stored products research 23(2): 119-123.

Cox, P., W. Parish, and M. Ledson. 1990. Factors Affecting the Refuge-Seeking Behavior of Cryptolestes ferrugineus (Stephens) (Coleoptera, Cucujidae). J. Stored Prod. Res. 26: 169-174.

Cox, P., J. Atkinson, K. Bannon, and J. Hatfield. 1993. Studies of Refuge-Seeking Behavior and Locomotory Activity in Cryptolestes ferrugineus (Stephens) (Coleoptera, Cucujidae) using Time-Lapse Video Recordings. J. Stored Prod. Res. 29: 283-289.

Duehl, Adrian J., Arbogast, R.T. and Teal, P.E. 2011a. Age and sex related responsiveness of Tribolium castaneum (Coleoptera: Tenebrionidae) in novel behavioral bioassays. Environmental Entomology 40: 82-87.

Duehl, A.J., Arbogast, R.T. and Teal, P.E. 2011b. Density-related volatile emissions and responses in the red flour beetle, Tribolium castaneum. Journal of Chemical Ecology 37: 525-532.

Fadamiro, H. Y. and Wyatt, T. D. 1996. Factors influencing response of flying Prostephanus truncatus to its male-produced aggregation pheromone. Physiol. Entomol. 21: 179-187.

Fardisi, M. and L. J. Mason. 2013. Influence of lure (food/sex pheromone) on young mated cigarette beetle (Lasioderma serricorne (F.)) (Coleoptera: Anobiidae) flight initiation. J. Stored Prod. Res. 53: 15-18.

Fardisi, M. and L. J. Mason. 2013. Influence of temperature, gender, age, and mating status on cigarette beetle (Lasioderma serricorne (F.)) (Coleoptera: Anobiidae) flight initiation. J. Stored Prod. Res. 52: 93-99.

Fontenot, Emily A., Frank H. Arthur and Kris L. Hartzer. “Oviposition of Dermestes maculatus DeGeer, the hide beetle, as affected by biological and environmental conditions.” Journal of Stored Products Research 64 (2015): 154-159.

Hagstrum, D. W., and Smittle, B. J. 1980. Age- and sex-specific tunneling rates of adult Tribolium castaneum. Ann. Entomol. Soc. Am. 73: 11-13.

Lextrait, P., Biemont, J-C. and Pouzat, J. 2008. Pheromone release by the two forms of Callosobruchus maculatus females: effects of age, temperature and host plant. Physiological Entomology 20: 309-317.

Lextrait, Patricia, Jean-Claude Biemont and Jean Pouzat. 1994. Comparison of walking locomotory reactions of two forms of Callosobruchus maculatus males subjected to female sex pheromone stimulation (Coleoptera: Bruchidae). Journal of chemical ecology 20(11): 2917-2930.

Mathieu F, Gaudichon,V., Brun, L.O. and Frerot, B., 2001. Effects of physiological status on olfactory and visual responses of female Hypothenemus hampei during host plant colonization. Physiol. Entomol. 26: 189–193.

Moriya, Seiichi and Satoshi Hiroyoshi. 1998. Flight and Locomotion Activity of the Sweet Potato Weevil (Coleoptera: Brentidae) in Relation to Adult Age, Mating Status, and Starvation. Journal of Economic Entomology 91 (2): 439-443.

Nguyen, D. T. 2008. Effects of starvation period on the locomotory response of Rhyzopertha dominica (F.). J. Stored Prod. Res. 44: 100-102.

ObengOfori, D. and T. Coaker. 1990. Some Factors Affecting Responses of 4 Stored Product Beetles (Coleoptera, Tenebrionidae and Bostrichidae) to Pheromones. Bull. Entomol. Res. 80: 433-441.

Perez-Mendoza, J., J. F. Campbell and J. E. Throne. 2011. Effects of Rearing Density, Age, Sex, and Food Deprivation on Flight Initiation of the Red Flour Beetle (Coleoptera: Tenebrionidae) J. Econ. Entomol. 104(2): 443-451.

Perez-Mendoza, J., J. F. Campbell and J. E. Throne. 2011. Influence of Age, Mating Status, Sex, Quantity of Food, and Long-Term Food Deprivation on Red Flour Beetle (Coleoptera: Tenebrionidae) Flight Initiation J. Econ. Entomol. 104(6): 2078-2086.

Perez-Mendoza, J., Dover, B. A., Hagstrum, D. W., and Baker, J. E. 1998. Flight activity of Rhyzopertha dominica (Coleoptera: Bostrichidae) in response to feeding damage and accumulation of waste. J. Econ. Entomol. 91: 1445-1448.

Perez-Mendoza, J., Dover, B. A., Hagstrum, D. W. and Hopkins, T. L. 1999. Effect of crowding, food deprivation, and diet on flight initiation and lipid reserves of the lesser grain borer, Rhyzopertha dominica. Entomol. Exp. Appl. 91: 317-326.

Perttunen, V. 1972 Humidity and light reactions of Sitophilus granarius (L.), S. oryzae L.(Col., Curculionidae), Rhizopertha dominica F.(Bostrychidae), and Acanthoscelides obtectus Say (Bruchidae).  Annales Entomologici Fennici 38: 161-176

Perttunen, V. and Hayrinen ,T. 1970. Effect of age on take-off activity in Acanthoscelides obtectus Say (Col., Bruchidae) Annales Entomologici Fennici 36: 35-39.

Perttunen, V. and K. M. Killstrom. 1971. Effect of desiccation on the light reactions of Calandra granaria L. and Calandra oryzae L. (Col., Curculionidae). Annales Entomologici Fennici 37: 147-154. (Sitophilus granarius, Sitophilus oryzae)

Perttunen, V. and M. Lahermaa. 1958. Reversal of negative phototaxis by desiccation in Tenebrio molitor L. (Col., Tenebrionidae). Annales Entomologici Fennici 24: 69-73.

Pierce, A., J. Borden, and A. Oehlschlager. 1983. Effects of Age and Population-Density on Response to Beetle and Food Volatiles by Oryzaephilus surinamensis and O. mercator (Coleoptera, Cucujidae). Environ. Entomol. 12: 1367-1374.

Rakowski, Grzegorz and B. Cymborowski. 1986. Some Environmental and Physiological Factors Influencing the Response of the Hide Beetle, Dermestes maculatus, to Aggregation Pheromone. International Journal of Invertebrate Reproduction and Development 9:35-41.

Rietdorf, K. and Steidle, J.L.M., 2002. Was Hopkins right? Influence of larval and early adult experience on the olfactory response in the granary weevil Sitophilus granarius (Coleoptera: Curculionidae). Physiological Entomology 27: 223-227.

Roth, L. M. and E. P. Willis . 1951. The effects of desiccation and starvation on the humidity behavior and water balance of Tribolium confusum and Tribolium castaneum. J. Exp. Zool. 118: 337–361.

Scholz, D., Borgemeister, C., Markham, R. H. and Poehling, H. M. 1997. Flight initiation in Prostephanus truncatus: Influence of population density and aggregation pheromone. Entomol. Exp. Appl. 85: 237-245.

Shimizu, T. and S. Moriya. 1996. Flight Time and Flight Age in the Sweetpotato Weevil, Cylas formicarius (Fabricius) (Coleoptera: Brentidae). Applied Entomology and Zoology 31: 575-580.

Surtees, G. 1965. Laboratory studies on dispersion behaviour of adult beetles in grain. IX. Some defferences in behaviour of mated and unmated grain weevils. Entomol. Mon. Mag. 100: 105-108.

Taylor, T. A., and Agbaje, L. A. 1974. Flight activity in normal and active forms of Callosobruchus maculatus (F.) in a store in Nigeria. J. Stored Prod. Res. 10:9-16.

Traynier, R. M . M . 1970. Sexual behaviour of the Mediterranean flour moth, Anagasta kuhniella: Some influences of age, photoperiod, and light intensity. Can. Entomol. 102: 534-540.

Utida, S. 1972. Density dependent polymorphism in the adult of Callosobruchus maculatus (Coleoptera: Bruchidae). J. Stored Prod. Res. 8: 111-126.

Ward, J.P. 1980. Post-Mating changes in the behaviour and activity of the female Anobiid bread beetle, Stegobium Paniceum. Physiological Enthomology 5: 301-306.

Ward J.P. 1981. Mating behavior and the mechanism of male orientation in the anobiid bread beetle, Stegobium Paniceum. Physiological Entomology 6: 213-217.

Weston, P. A. and Hoffman, S. A. 1992. Influence of starvation, dehydration and humidity differential on humidity response of Sitophilus zeamais (Coleoptera: Curculionidae). Environ. Entomol. 21: 1345-1350.

Wright, E. J., and Morton, R. 1995. Daily flight activity of Trogoderma variabile (Coleoptera: Dermestidae) and Rhyzopertha dominica (Coleoptera: Bostrichidae). J. Stored Prod. Res. 31:177-184.

Gradient orientation

Amos, T. G. 1968. Some observations on Oryzaephilus surinamensis L. (Coleoptera: Silvanidae) in a temperature-moisture gradient. Entomologist 101: 67-70.

Amos, T. G. 1969. Reactions of Carpophilus spp. (Coleoptera, Nitidulidae) to humidity with reference to gradient steepness. Anim. Behav. 17: 9-13.

Amos, T. G. and F. L. Waterhouse. 1967. Phasic behaviour shown by two Carpophilus species (Coleoptera, Nitidulidae) in various humidity gradients and its ecological significance. Oikos 18: 345-50.

Amos, T. G. and F. L. Waterhouse. 1969. The distribution and biology of tribolium castaneum (Coleoptera, Tenebrionidae) on temperature gradients varying in steepness. Entomol. Exp. Appl. 12: 44-52.

Amos, T. G., F. L. Waterhouse and N. A. Chetham. 1968. Temporal Distribution of Tribolium castaneum Herbst and Cadra cautella Walker on Temperature Gradients. Separatum Esperientia 24: 86-87.

Amos, T. G., F. L. Waterhouse and N. A. Dicker. 1969. Preliminary observations on the distribution and biology of Tribolium castaneum on a temperature gradient. Tribolium Information Bulletin 11: 67-68.

Amos, T., and S. Dennler. 1969. A Comparison of 3 Strains of Oryzaephilus surinamensis (L) (Col, Silvanidae) on a Temperature-Mositure Gradient. J. Stored Prod. Res. 5: 173-175.

Bell, C. H. 1986. The response of insects to fumigant concentration gradients. p. 316-326 In E. Donahaye and S. Navarro eds. Proc. 4th Intern. Work. Conf. Stored-Prod. Prot. Wallach Press, Jerusalem.

Collins, L. E. and S. T. Conyers. 2009. Moisture content gradient and ventilation in stored wheat affect movement and distribution of Oryzaephilus surinamensis and have implications for pest monitoring. J. Stored Prod. Res. 45: 32-39.

Deal, John. 1941. The temperature preferendum of certain insects. J. Animal Ecology 10: 323-356. (Acanthoscelides obtectus, Anthrenus verbasci, Cryptolestes turcicus, Dermestes, Gnathocerus cornutus, Lasioderma serricorne, Oryzaephilus mercator, Oryzaephilus surinamensis, Ptinus tectus, Sitophilus granarius, Sitophilus oryzae, Sitotroga cerealella, Stegobium paniceum, Tenebrio molitor, Tribolium confusum, Tineola bisselliella)

Dicker, N. A. and Waterhouse, F. L. 1972. Life history distributions of geographically different populations of Ephestia cautella (Walker) (Lep., Phycitidae) in gradient environments. Norsk Entomologisk Tidsskrift 19: 11-15.

Fletcher, L. W., W. Knulle, D. P. Childs, R. R. Spadafora, J. S. Long and C. D. Delamar. 1973. Responses of cigarette beetle larvae to temperature and humidity. Agricultural Research Service, Southern Region, USDA, ARS-S-22. (temperature and humidity gradients)

Flinn, P. and D. Hagstrum. 1998. Distribution of Cryptolestes ferrugineus (Coleoptera : Cucujidae) in response to temperature gradients in stored wheat. J. Stored Prod. Res. 34: 107-112.

Flinn, P. W. and D. W. Hagstrum. 2011. Movement of Rhyzopertha dominica in response to temperature gradients in stored wheat. J. Stored Prod. Res. 47: 407-410.

Graham, W. W. 1965. Biology and behaviour studies in a temperature gradient., p. 345-346. In Proceedings of the 12th International Congress Entomology, London, 8-16 July 1964. Royal Entomological Society of London.

Graham, W. M., and F. L. Waterhouse. 1964. The distribution and orientation of Tribolium on inclines, and the concept of controls in gradient experiments. Animal Behaviour 12(2): 368-373.

Hagstrum, D., P. Flinn and J. Gaffney. 1998. Temperature gradient on Tribolium castaneum (Coleoptera : Tenebrionidae) adult dispersal in stored wheat. Environ. Entomol. 27: 123-129.

Hubert, J., S. Pekár, M. Nesvorná and V. Šustr 2010. Temperature Preference and Respiration of Acaridid Mites.  Journal of Economic Entomology 103(6): 2249–2257. (Acarus siro, Dermatophagoides farinae, Lepidoglyphus destructor, Tyrophagus putrescentiae)

Jian, F., D. Jayas and N. White. 2003. Movement of adult rusty grain beetles, Cryptolestes ferrugineus (Coleoptera : Cucujidae), in wheat in response to 5 degrees C/m temperature gradients at cool temperatures. J. Stored Prod. Res. 39: 87-101.

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