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United States Department of Agriculture

Agricultural Research Service

John A. Byers

Research Entomologist


Pest Management & Biological Control Research Unit

USDA-ARS, Arid-Land Agricultural Research Center


John A. Byers

Research Entomologist

Arid-Land Agricultural Research Center

21881 North Cardon Lane

Maricopa, Arizona, USA 85138


520-316-6330 (FAX)



B.S. Entomology, Colorado State University, 1971

M.S. Entomology, Colorado State University, 1973

Ph.D. Entomology, University of California at Berkeley, 1978

Research Interests:
Models of insect mate finding, host plant finding and host selection, dispersal, predator-prey encounter rates, mass trapping, and behavioral interactions.
Chemical ecology of plant-insect interactions and plant resistance
Isolation and identification of pheromones, host plant attractants, non-host deterrents and repellents, and other natural semiochemicals
Electronic sensors and data loggers for continuous measurements of environmental parameters.
Chemical ecology of bark beetles and conifers, semiochemicals of fire ants, and evolution and structures of moth pheromone components. (earlier work)
New interests include whiteflies, plant bugs (Lygus), parasitoids, pink bollworm, aphids, lacewings, plant volatiles, and ant predators of cotton insect pests. I am also involved in GC-MS and GC-EAD methods. (present studies)

Key Publications

2010 - 2012

Byers, J.A. 2012a. A population genetic model of evolution of host-mate attraction and nonhost repulsion in a bark beetle Pityogenes bidentatus. Psyche: A Journal of Entomology vol. 2012, ID 529573, pp. 1-9. [PDF]

Byers, J.A. 2012b. Estimating insect flight densities from attractive trap catches and flight height distributions. Journal of Chemical Ecology 38:592-601. [PDF]

Byers, J.A. 2012c. Bark beetles, Pityogenes bidentatus, orienting to aggregation pheromone avoid conifer monoterpene odors when flying but not when walking. Psyche: A Journal of Entomology vol. 2012, ID 940962, pp. 1-10. [PDF]

Byers, J.A. 2012d. Modelling female mating success during mass trapping and natural competitive attraction of searching males or females. Entomologia Experimentalis et Applicata 145:228-237. [PDF]

Byers, J.A. 2012e. Ecological interactions of bark beetles with host trees.
Psyche: A Journal of Entomology vol. 2012, ID 252961, pp. 1-3. [PDF]

Byers, J.A. and G. Birgersson. 2012. Host-tree monoterpenes and biosynthesis of aggregation pheromones in the bark beetle Ips paraconfusus. Psyche: A Journal of Entomology vol. 2012, ID 539624, pp. 1-10. [PDF]

Han, B., Zhang, Q.H., and Byers, J.A. 2012. Attraction of the tea aphid, Toxoptera aurantii, to combinations of volatiles and colors related to tea plants. Entomologia Experimentalis et Applicata 144:258-269. [PDF]

Lu, R.C., Wang, H.B., Zhang, Z., Byers, J.A., Jin, Y.J., Wen, H.F., Shi, W.J. 2012. Coexistence and competition between Tomicus yunnanensis and T. minor (Coleoptera: Scolytinae) in Yunnan pine. Psyche: A Journal of Entomology vol. 2012, ID 185312, pp. 1-6. [PDF]

Lu, R.C., Wang, H.B., Zhang, Z., Byers, J.A., Jin, Y.J., Wen, H.F., Shi, W.J. 2012. Attraction of Tomicus yunnanensis (Coleoptera: Scolytinae) to Yunnan pine logs with and without periderm or phloem: an effective monitoring bait. Psyche: A Journal of Entomology vol. 2012, ID 794683, pp. 1-5. [PDF]

Rodriguez-Saona, C.R., Byers, J.A., and Schiffhauer, D. 2012. Effect of trap color
and height on captures of blunt-nosed and sharp-nosed leafhoppers (Hemiptera:
Cicadellidae) and non-target arthropods in cranberry bogs. Crop Protection 40:132-144. [PDF]

Brent, C.S. and Byers, J.A. 2011. Female attractiveness modulated by a male-derived antiaphrodisiac pheromone in a plant bug. Animal Behaviour 82:937-943.[PDF]

Byers, J.A. 2011. Analysis of vertical distributions and effective flight layers of insects: Three-dimensional simulation of flying insects and catch at trap heights. Environmental Entomology 40:1210-1222.[PDF]

Byers, J.A. and Levi-Zada, A. 2011. Individual variation of (S)-4-methyl-3-heptanone in heads of braconid wasp, Leiophron uniformis, and Pogonomyrmex ants indicates costs of semiochemical production. Chemoecology 21:35-44.[PDF]

Zhang, Q.H., Ma, J.H., Yang, Q.Q., Byers, J.A., Klein, M. G., Zhao, F.Y., and Luo, Y.Q. 2011. Olfactory and visual responses of the long-legged chafer Hoplia spectabilis Medvedev (Coleoptera: Scarabaeidae) in Qinghai province, China. Pest Management Science 67:162-169.[PDF]

Salvucci, M.E., Barta, C., Byers, J.A., and Canarini, A. 2010. Photosynthesis and assimilate partitioning between carbohydrates and isoprenoid products in vegetatively active and dormant guayule: physiological and environmental constraints on rubber accumulation in a semi-arid shrub. Physiologia Plantarum 140:368-379.[PDF]

Byers, J.A., and Zhang, Q.H. 2010. Chemical Ecology of Bark Beetles in regard to search and selection of host trees, pp. 88-111. In, Recent Advances in Entomological Research (eds.) T.-X. Liu and L. Kang. The Higher Education Press, Beijing, China.

2005 - 2009

Huang, C. H., Yan, F. M., Byers, J. A., Wang, R. J., and Xu, C. R. 2009. Volatiles induced by the larvae of the Asian corn borer (Ostrinia furnacalis) in maize plants affect behavior of conspecific larvae and female adults. Insect Science 16:311-320. [PDF]

El-Sayed, A. M, Suckling, D. M., Byers, J. A., Jang, E. B., and Wearing, C. H. 2009. Potential of ‘lure and kill’ in long-term pest management and eradication of invasive species. Journal of Economic Entomology 102:815-835. [PDF]
Zada, A., Falach, L., and Byers, J. A. 2009. Development of sol-gel formulations for slow release of pheromones. Chemoecology 19:37-45. [PDF]

Byers, J. A. 2009. Modeling distributions of flying insects: Effective attraction radius of pheromone in two and three dimensions. Journal of Theoretical Biology 256:81-89. [PDF]

Blackmer, J. L., and Byers, J.A. 2009. Lygus spp. (Heteroptera: Miridae) host-plant interactions with Lesquerella fendleri (Brassicaceae), a new crop in the arid southwest. Environmental Entomology 38:159-167. [PDF]

Byers, J. A. 2008. Active space of pheromone plume and its relationship to effective attraction radius in applied models. Journal of Chemical Ecology 34:1134-1145. [PDF]

El-Sayed, A. M, Byers, J. A., Manning, L. M., Jürgens, A., Mitchell, V. J., and Suckling, D. M. 2008. Floral scent of Canada thistle and its potential as a generic insect attractant. Journal of Economic Entomology 101:720-727. [PDF]

Blackmer, J. L., Byers, J. A., and Rodriguez-Saona, C. 2008. Evaluation of color traps for monitoring Lygus spp.: Design, placement, height, time of day, and non-target effects. Crop Protection 27:171-181. [PDF]

Natwick, E. T, Byers, J. A., Chu, C. C., Lopez, M., and Henneberry, T. J. 2007. Early detection and mass trapping of Frankliniella occidentalisand Thrips tabaciin vegetable crops. Southwestern Entomologist 32:229-238. [PDF]

Byers, J. A. 2007. Simulation of mating disruption and mass trapping with competitive attraction and camouflage. Environmental Entomology 36:1328-1338. [PDF] 

Byers, J. A. 2006. Production and predator-induced release of volatile chemicals by the plant bug Lygus hesperus. Journal of Chemical Ecology 32:2205-2218. [PDF]

El-Sayed, A. M, Suckling, D. M, Wearing, C. H., and Byers, J. A. 2006. Potential of mass trapping for long-term pest management and eradication of invasive species. Journal of Economic Entomology 99:1550-1564. [PDF]

Byers, J. A. 2006. Analysis of insect and plant colors in digital images using Java software on the Internet. Annals of the Entomological Society of America 99:865-874. [PDF]

Thoss, V., and Byers, J. A. 2006. Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization. Chemoecology 16:51-58. [PDF]

Byers, J. A. 2006. Pheromone component patterns of moth evolution revealed by computer analysis of the Pherolist. Journal of Animal Ecology 75:399-407. [PDF]

Byers, J. A., and Castle, S. J. 2005. Areawide models comparing synchronous versus asynchronous treatments for control of dispersing insect pests. Journal of Economic Entomology 98:1763-1773. [PDF]

Byers, J. A. 2005. Chemical constraints on the evolution of olfactory communication channels of moths. Journal of Theoretical Biology 235:199-206. [PDF]

Byers, J. A. 2005. A cost of alarm pheromone production in cotton aphids, Aphis gossypii. Naturwissenschaften 92:69-72. [PDF]

2000 - 2004

Blackmer, J. L., Rodriquez-Saona, C., Byers, J. A., Shope, K. L., and Smith, J. P. 2004. Behavioral response of Lygus hesperus to conspecifics and headspace volatiles of alfalfa in a Y-tube olfactometer. Journal of Chemical Ecology 30:1547-1564. [PDF]

Byers, J. A. 2004. Equations for nickel-chromium wire heaters of column transfer lines in gas chromatographic-electroantennographic detection (GC-EAD). Journal of Neuroscience Methods 135:89-93. [PDF]

Byers, J. A., Zhang, Q. H., and Birgersson, G. 2004. Avoidance of nonhost plants by a bark beetle, Pityogenes bidentatus, in a forest of odors. Naturwissenschaften 91:215-219. [PDF]

Byers, J. A. 2004. Chemical ecology of bark beetles in a complex olfactory landscape. pp. 89-134. In, Bark and Wood Boring Insects in Living Trees in Europe, a Synthesis, F. Lieutier, K. R. Day, A. Battisti, J. C. Grégoire, H. Evans (Eds). Kluwer Academic Publishers, Dordrecht, The Netherlands. [PDF]

Byers, J. A. 2002. Internet programs for drawing moth pheromone analogs and searching literature database. Journal of Chemical Ecology 28:807-817. [PDF]

Byers, J. A. 2001. Correlated random walk equations of animal dispersal resolved by simulation. Ecology 82:1680-1690. [PDF]

Schlyter F., Zhang Q-H., Anderson, P., Byers, J. A., Wadhams, L. J., Löfqvist, J., and Birgersson, G. 2000. Electrophysiological and behavioural responses of Scandinavian Tomicus piniperdaand T. minor(Coleoptera: Scolytidae), to non-host leaf and bark volatiles. Canadian Entomologist 132:965-981. [PDF]

El-Sayed, A. M. and Byers, J. A. 2000. Inhibitory effect of monoterpenes on response of Pityogenes bidentatusto aggregation pheromone released by piezoelectric sprayer for precision release of semiochemicals. Journal of Chemical Ecology 26:1795-1809. [PDF]

Byers, J. A., Zhang, Q. H., and Birgersson, G. 2000. Strategies of a bark beetle, Pityogenes bidentatus, in an olfactory landscape. Naturwissenschaften 87:503-507. [PDF] 

Byers, J. A. 2000. Wind-aided dispersal of simulated bark beetles flying through forests. Ecological Modelling 125:231-243. [PDF]

Byers, J. A. 2000. Web Site: Chemical Ecology of Insects( Reviewed by Kaiser, J. 2000. "Bug Juice". Science 288:767. (web site now at:

1995 - 1999

Byers, J. A. 1999. Effects of attraction radius and flight paths on catch of scolytid beetles dispersing outward through rings of pheromone traps. Journal of Chemical Ecology 25:985-1005. [PDF]

Byers, J. A. 1999. Interactive learning using expert system quizzes on the internet. Educational Media International 36:191-194. [PDF]

Byers, J. A., Zhang, Q. H., Schlyter, F., and Birgersson, G. 1998. Volatiles from nonhost birch trees inhibit pheromone response in spruce bark beetles. Naturwissenschaften 85:557-561. [PDF]

Byers, J. A. 1996. Random selection algorithms for spatial and temporal sampling. Computers in Biology and Medicine 26:41-52. [PDF]

Byers, J. A. 1996. An encounter rate model of bark beetle populations searching at random for susceptible host trees. Ecological Modelling 91:57-66. [PDF]

Byers, J. A. 1996. Taxokey: a universal taxonomic key program using DOS text files and graphics. Computer Applications in the Biosciences 12:185-189. [PDF]

Byers, J. A. 1996. Correct calculation of Dirichlet polygon areas. Journal of Animal Ecology 65:528-529. [PDF]

Byers, J. A. 1996. Temporal clumping of bark beetle arrival at pheromone traps: Modeling anemotaxis in chaotic plumes. Journal of Chemical Ecology 22:2133-2155. [PDF]

Byers, J. A. 1995. Host tree chemistry affecting colonization in bark beetles, in R.T. Cardé and W.J. Bell (eds.). Chemical Ecology of Insects 2. Chapman and Hall, New York, pp. 154-213. [PDF]

Zolubas, P. and Byers, J. A., 1995. Recapture of dispersing bark beetles Ips typographusL. (Col., Scolytidae) in pheromone-baited traps. Journal of Applied Entomology 19:285-289. [PDF]

1990 - 1994

Byers, J. A. 1993. Randomization algorithms in BASIC for experimental design. Computers in Biology and Medicine 23:167-176. [PDF]

Byers, J. A. 1993. Avoidance of competition by spruce bark beetles, Ips typographusand Pityogenes chalcographus. Experientia 49:272-275. [PDF]

Byers, J. A. 1993. Simulation and equation models of insect population control by pheromone-baited traps. Journal of Chemical Ecology 19:1939-1956. [PDF]

Byers, J. A. 1993. Orientation of bark beetles Pityogenes chalcographusand Ips typographusto pheromone-baited puddle traps placed in grids: A new trap for control of scolytids. Journal of Chemical Ecology 19:2297-2316. [PDF]

Byers, J. A. 1993. Conversion of BIOSIS Previews database files to reference text files and reprint request forms. American Entomologist 39:245-248. [PDF]

Zhang, Q. H., Byers, J. A., and Zhang, X. D. 1993. Influence of bark thickness, trunk diameter and height on reproduction of the long- horned beetle, Monochamus sutor(Col., Cerambycidae) in burned larch and pine. Journal of Applied Entomology 115:145-154. [PDF]

Byers, J. A. 1992. Grid cell contour mapping of point densities: bark beetle attacks, fallen pine shoots, and infested trees. Oikos 63:233-243. [PDF]

Byers, J. A. 1992. Ref-list: Program to list references found in DOS text files of scientific manuscripts. Computer Applications in the Biosciences 8:196-198. [PDF]

Byers, J. A. 1992. Dirichlet tessellation of bark beetle spatial attack points. Journal of Animal Ecology 61:759-768. [PDF]

Byers, J. A. 1992. Trends in chemical ecology revealed with a personal computer program for searching data bases of scientific references and abstracts. Journal of Chemical Ecology 18:1481-1495. [PDF]

Byers, J. A. 1992. Optimal fractionation and bioassay plans for isolation of synergistic chemicals: the subtractive-combination method. Journal of Chemical Ecology 18:1603-1621. [PDF]

Byers, J. A. 1992. Attraction of bark beetles, Tomicus piniperda, Hylurgops palliatus, and Trypodendron domesticum and other insects to short-chain alcohols and monoterpenes. Journal of Chemical Ecology 18:2385-2402. [PDF]

Schlyter, F., Birgersson, G., Byers, J. A., and Bakke, A. 1992. The aggregation pheromone of Ips duplicatus and its role in competitive interactions with I. typographus(Coleoptera: Scolytidae). Chemoecology 3:103-112. [PDF]

Zhang, Q. H., Byers, J. A., and Schlyter, F. 1992. Optimal attack density in the larch bark beetle, Ips cembrae(Coleoptera: Scolytidae). Journal of Applied Ecology 29:672-678. [PDF]

Byers, J. A. 1991. Simulation of mate-finding behaviour of pine shoot beetles, Tomicus piniperda.Animal Behaviour 41:649-660. [PDF]

Byers, J. A. 1991. BASIC algorithms for random sampling and treatment randomization. Computers in Biology and Medicine 21:69-77. [PDF]

Byers, J. A. 1991. Cardcat: A scientific reference manager for IBM- compatible personal computers. American Entomologist 37:240-242. [PDF]

Byers, J. A. 1991. Pheromones and chemical ecology of locusts. Biological Reviews 66:347-378. [PDF]

Birgersson, G., Byers, J. A., Bergström, G., and Löfqvist, J. 1990. Production of pheromone components, chalcogran and methyl (E,Z)-2,4- decadienoate, in the spruce engraver Pityogenes chalcographus. Journal of Insect Physiology 36:391-395. [PDF]

Byers, J. A., and Birgersson, G. 1990. Pheromone production in a bark beetle independent of myrcene precursor in host pine species. Naturwissenschaften 77:385-387. [PDF]

Byers, J. A., Birgersson, G., Löfqvist, J., Appelgren, M., and Bergström, G. 1990. Isolation of pheromone synergists of bark beetle, Pityogenes chalcographus, from complex insect-plant odors by fractionation and subtractive-combination bioassay. Journal of Chemical Ecology 16:861-876. [PDF]

Byers, J. A., Schlyter, F., Birgersson, G., and Francke, W. 1990. E-Myrcenol in Ips duplicatus: An aggregation pheromone component new for bark beetles. Experientia 46:1209-1211. [PDF]

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Last Modified: 3/21/2013