Department of Entomology


Ring Cardéucr entomology

Distinguished Professor of Entomology and
A.M. Boyce Endowed Chair
Location: Boyden 216
Tel: (951) 827-4492
E-mail: ring.carde@ucr.edu





Former Institution

University of Massachusetts, Amherst (1981-1996) and Michigan State University (1975-1981)


BS Biology 1966
Tufts University
MS Entomology 1968
Cornell University
PhD Entomology 1971
Cornell University


Certificate of Achievement for Outstanding Achievements, International Congress of Entomology, 2016.
President, International Society of Chemical Ecology, 2012-2013
Visiting Professor, Swedish Agricultural University (SLU) Alnarp, 2011
Silver Medal, International Society of Chemical Ecology, 2009
Elected Fellow, Entomological Society of America, 1998
Elected Fellow, American Association for the Advancement of Science, 1997
Elected Fellow, Royal Entomological Society, 1992
Elected Fellow, Entomological Society of Canada, 1992
Chancellor's Medal, University of Massachusetts, 1988
Distinguished Faculty Lecturer, University of Massachusetts, 1988
Visiting Scientist, Wageningen University, 1988
Recognition Award for Outstanding Contributions to Agriculture, Entomological Society of America, 1980

Research Area

Insects employ a variety of chemical cues to find and identify resources. We study two model systems, moth communication by pheromones and orientation by female mosquitoes to odors from prospective hosts. Some of our work with moths is aimed at understanding mechanisms and evolutionary forces mediating these odor-mediated behaviors, and includes documenting within species variation in pheromone production and response, and using measurements of heritability and selection experiments to determine the genetic architecture of pheromone production and response. With mosquitoes our efforts are directed at understanding how host-emitted odors influence orientation from a distance and landing on the host, and how repellents interfere with these behaviors. Our work on odor-mediated orientation includes investigation of dispersal of odor plumes in wind, the role of plume structure in orientation, and the integration of visual and odor inputs in modulating orientation. Many of these studies employ wind tunnels where we can manipulate the sensory inputs and record maneuvers with 3-D video. We also study the use of formulated synthetic pheromone to disrupt mating of moth pests.



Bau, J. and R.T. Cardé. 2016. Simulation modeling to interpret the captures of moths in pheromone-baited traps used for surveillance of invasive species: the gypsy moth as a model case. J. Chem. Ecol. 42:877-887.

Cardé, R.T. 2016. Moth Navigation along Pheromone Plumes. In: Pheromone Communication in Moths: Evolution, Behavior and Application. J.D. Allison and R.T. Cardé (eds.).  University of California Press, Berkeley, pp. 173-189.

Allison, J.D. and R.T. Cardé. 2016. Variation in Moth Pheromones: Causes and Consequences. In: Pheromone Communication in Moths: Evolution, Behavior and Application. J.D. Allison and R.T. Cardé (eds.). University of California Press, Berkeley, pp. 11-23.

Allison, J.D. and R.T. Cardé. 2016. Pheromones: Reproductive Isolation and Evolution in Moths. In: Pheromone Communication in Moths: Evolution, Behavior and Application. J.D. Allison and R.T. Cardé (eds.).  University of California Press, Berkeley, pp. 25-41.

Choi, M.-Y., S.-J. Ahn, K.-C. Park, R. Vander Meer, R. T. Cardé and R. Jurenka. 2016. Heliothine moths contain aldehydes and butyrate esters as potential pheromones. J. Chem. Ecol. J. Chem. Ecol.  42:425-432.

Webster, B. and R. T. Cardé. 2016. Use of habitat odour by host-seeking insects. 2016. Biol. Reviews. DOI 10.1111/brv.12281.

Bau, P. and R.T. Cardé. 2015. Modeling optimal strategies for finding a resource-linked, windborne odor plume: theories, robotics and biomimetic lessons from flying insects. Integr. Comp. Biol. 55:461-477.

Cardé, R.T. 2015. Multi-cue integration: how mosquitoes find a human host. Curr. Biol. 25:R793-795.

Wheeler, C.A., J.G. Millar and R.T. Cardé. 2015. Multimodal signal interactions in the ladybeetle, Hippodamia convergens, aposematic system. Chemoecology 25:123-133.

Webster, B., E.S. Lacey and R.T. Cardé. 2015. Waiting with bated breath: opportunistic orientation to human odor in the malaria mosquito, Anopheles gambiae, is modulated by minute changes in carbon dioxide concentration. J. Chem. Ecol. 41:59-66.

Cardé, R.T. 2014. Defining attraction and aggregation pheromones: teleological versus functional perspectives. J. Chem. Ecol. 40: 519-520.

Wheeler, C.A. and R.T. Cardé. 2014. Following in their footsteps: cuticular hydrocarbons as overwintering aggregation site markers in Hippodamia convergens. J. Chem. Ecol. 40:418-428.

Collett, M. and R.T. Cardé. 2014. Insect navigation: many senses make efficient foraging paths.  Curr. Biol. 24:R362-364.

Lacey, E.S., A. Ray and R.T. Cardé. 2014. Close encounters: contributions of carbon dioxide and human skin odour to finding and landing on a host in female Aedes aegypti. Physiol. Entomol. 39:60-68.

Girling, R.D., B.S. Higbee and R.T. Cardé. 2013. The plume also rises: trajectories of pheromone plumes issuing from point sources in an orchard canopy at night. J. Chem. Ecol. 39:1150-1160.

Wheeler, C.A. and R.T. Cardé. 2013. Defensive allomones function as aggregation pheromones in diapausing ladybird beetles, Hippodamia convergens. J. Chem. Ecol. 39:723-732.

Kárpáti, Z., M. Tasin, R.T. Cardé and T. Dekker. 2013. Early quality assessment lessens pheromone specificity in a moth. Proc. Nat. Acad. Sci. USA. 110:7377-7382.

Lacey, E.S. and R.T. Cardé. 2012. Location of and landing on a source of human body odour by female Culex quinquefasciatus in still and moving air Physiol. Entomol. 37:153-159.

Wisniewska, J. and R. T. Cardé. 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

Cardé, R.T., A.M. Cardé and R.D. Girling. 2012. Observations on the flight paths of the day-flying moth Virbia lamae during periods of mate location: Do males have a strategy for contacting the pheromone plume? J. Anim. Ecol. 81:268-276

Dekker, T. and R.T. Cardé. 2011. Moment-to-moment flight manoeuvres of the female yellow fever mosquito (Aedes aegypti L.) in response to plumes of carbon dioxide and human skin odour. J. Exp. Biol. 214:3480-3494.

Turner, S.L., N. Li, T. Guda, J. Guthrie, R.T. Cardé and A. Ray. 2011. Ultra-prolonged activation of CO2-sensing neurons disorients mosquitoes. Nature 474:87-91.

Lacey, E.S. and R.T. Cardé. 2011. Activation, orientation, and landing of female Culex quinquefasciatus in response to carbon dioxide and odour from human feet; 3-D flight analysis in a wind tunnel. Med. Vet. Entomol. 25:94-103.

Cardé, R. T. and G. Gibson. 2010. Long-distance orientation of mosquitoes to host odours and other host-related cues. In: Ecology of Vector-Borne Diseases. Vol. 2. Olfaction in Vector-Host Interactions. W. Takken and B.G.F. Knols (eds.) Wageningen Academic Publishers. Wageningen, The Netherlands, pp. 115-141.

Kanno, H., L.P.S. Kuenen, K.A. Klingler, J.G. Millar and R.T. Cardé. 2010. Attractiveness of a four-component pheromone blend to male navel orangeworm moths. J. Chem. Ecol. 36:584-591.

Wang, H.-L., C.-H. Zhao, J. G. Millar, R.T. Cardé and C. Löfstedt. 2010. Biosynthesis of unusual moth pheromone components involves two distinctly different pathways in the navel orangeworm, Amyelois transitella. J. Chem. Ecol. 36:535-547.

Cardé, R.T. 2009. Orientation. In: Encyclopedia of Insects, 2nd edn. V.H. Resh and R.T. Cardé (eds.), Academic Press, San Diego, pp. 729-732.

Cardé, R.T. and J.G. Millar. 2009. Pheromones. In: Encyclopedia of Insects, 2nd edn. V.H. Resh and R.T. Cardé (eds.), Academic Press, San Diego pp. 766-772.

Koch, U.T, W. Lüder, U. Andrick, R.T. Staten and R.T. Cardé. 2009. Measurement by electroantennogram of airborne pheromone in cotton treated for mating disruption of Pectinophora gossypiella following removal of pheromone dispensers. Entomol. Exp. Appl. 130:1-9.

Zaspel, J.M., S.J. Weller and R.T. Cardé. 2008. A review of Virbia (formerly Holomelina) of America north of Mexico (Arctiidae: Arctiinae: Arctiini). Bull. Florida Mus. Nat. Hist. 48:59-113.

Cardé, R.T. 2008 Animal migration: seasonal reversals of migrant moths. Curr. Biol. 18: R1007-R1009.

Cardé, R.T. and J.G. Millar. 2008. The scent of a female: sex pheromones of female tiger moths. In: Tiger Moths and Wooly Bears: Behavior, Ecology and Natural History of the Arctiidae, W.E. Conner (ed.), Oxford University Press, New York. pp. 127-143.

Allison, J.D., D.A. Roff and R.T. Cardé. 2008. Genetic independence of female signal form and male receiver design in the almond moth, Cadra cautella. J. Evol. Biol. 21:1666-1672.

Cooperband, M.F., J.S. McElfresh, J.G. Millar and R.T. Cardé. 2008. Attraction of Culex quinquefasciatus Say (Diptera: Culicidae) to odors from chicken feces. J. Insect Physiol. 53:1184-1192.

Cardé, R.T. and M.A. Willis. 2008. Navigational strategies used by flying insects to find distant, wind-borne sources of odor. J. Chem. Ecol. 43:854-866

Cardé, R.T. 2008. High flying migrant moths: do they know where they are heading? Curr. Biol. 18:R472-R424.

Allison, J.D. and R.T. Cardé. 2008. Male pheromone blend preference function measured in choice and no-choice wind tunnel trials with almond moths, Cadra cautella. Anim. Behav. 75:259-266.

Zahn, D.K., R.D. Girling, S.J. McElfresh, R.T. Cardé and J.G. Millar. 2008. Biology and reproductive behavior of Murgantia histrionica (Heteroptera: Pentatomidae). Ann. Entomol. Soc. Amer. 101:215-228.

Allison, J.D. and R.T. Cardé. 2007. Bidirectional selection for novel pheromone blend ratios in the almond moth, Cadra cautella. J. Chem. Ecol. 33:2293-2307.

Girling, R.D. and R.T. Cardé. 2007. Analysis and manipulation of odor plumes from a piezo-electric release system and measurements of upwind flight of male almond moths, Cadra cautella, to pheromone plumes. J. Chem. Ecol. 33:1927-1945.

Braks, M.A.H. and R.T. Cardé. 2007. Improving efficacy of box gravid traps for collecting Culex quinquefasciatus. J. Vector Ecol. 32:83-89.

Braks, M.A.H., W.S. Leal and R.T. Cardé. 2007. Oviposition responses of gravid female Culex quinquefasciatus to egg rafts and low doses of oviposition pheromone under semifield conditions. J. Chem. Ecol. 33:567-578.

Cardé, R.T. 2007. Using pheromones to disrupt mating of moth pests. In: Perspectives in Ecological Theory and Integrated Pest Management, M. Kogan and P. Jepson (eds.) Cambridge University Press, Cambridge. pp. 122-169.

Allison, J.D. and R.T. Cardé. 2006. Heritable variation in the sex pheromone of the almond moth, Cadra cautella. J. Chem Ecol. 32:621-641.

Girling, R.D. and R. T. Carde. 2006. Analysis of the courtship behavior of the navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae), with a commentary on methods for the analysis of sequences of behavioral transitions. J. Insect Behav. 19:497-520.

Vetter, R.S., A.M. Sage, K.A. Justus, R.T. Cardé and C.G. Galizia. 2006. Temporal integrity of an airborne odor stimulus is greatly affected by physical aspects of the odor delivery system. Chem Senses. 31:359-369.

Cooperband, M.F. and R.T. Cardé. 2006. Orientation of Culex mosquitoes to carbon dioxide-baited traps: flight manoeuvres and trapping efficiency. Med. Vet. Entomol. 26:11-26.

Cooperband, M.F. and R.T. Cardé. 2006. Comparison of plume structures of carbon dioxide emitted from different mosquito traps. Med. Vet. Entomol. 26:1-10.

Braks, M.A.H., M.F. Cooperband and R.T. Cardé. 2005. 3-D flight track analyses of Culex mosquito orientation to oviposition-related odors. Proceedings Measuring Behavior 2005, Wageningen, The Netherlands, pp. 209-210.

Cardé, R.T. 2005. Bioassays for insect attractants, to what end: aiding chemical identification, measuring adaptive value or verifying orientation mechanisms? Proceedings Measuring Behavior 2005, Wageningen, The Netherlands, pp. 203-206.

Bau, J., K.A. Justus, C. Loudon and R.T. Cardé. 2005. Electroantennographic resolution of pulsed pheromone plumes in two species of moths with bipectinate antennae. Chem. Senses. 30: 771-780.

Dekker, T., M. Geier and R.T. Cardé. 2005. Carbon dioxide instantly sensitizes female yellow fever mosquitoes to human skin odours. J. Exp. Biol. 208:2963-2972.

Owen, J.P., B.A. Mullens, K.A. Justus and R.T. Cardé. 2005. Northern fowl mite orientation in a thermal gradient and evidence for idiothetic course control. Physiol. Entomol. 30:293-302.

Justus, K.A., R.T. Cardé and A.S. French. 2005. Dynamic properties of antennal responses in two moth species. J. Neurophysiol. 93:2233-2239.

Schofield, S.W., K.A. Justus, A. Mafra-Neto and R.T. Cardé. 2003. Flight of male Cadra cautella along plumes of air and pheromone superimposed on backgrounds of pheromone. Entomol. Exp. Appl. 109:173-182.

Koch, U., A.M. Cardé and R.T. Cardé. 2002. Calibration of an EAG system to measure airborne concentration of pheromone formulated for mating disruption of the pink bollworm moth, Pectinophoro gossypiella (Saunders) (Lep., Gelechiidae) J. Appl. Entomol. 126:431-436.

Dekker, T., B. Steib, R.T. Cardé and M. Geier. 2002. L-Lactic acid: a human-signifying host cue for the anthropophilic mosquito Anopheles gambiae. Med. Vet. Entomol. 16:91-98.

Justus, K.A., S.W. Schofield, J. Murlis and R.T. Cardé. 2002. Flight behaviour of Cadra cautella males in rapidly pulsed pheromone plumes. Physiol. Entomol. 27:58-66.

Justus, K.A. and R. T. Cardé. 2002. Flight behaviour of two moths, Cadra cautella and Pectinophora gossypiella, in homogeneous clouds of pheromone. Physiol. Entomol. 27:67-75.

Bau, J., K.A. Justus and R.T. Cardé. 2002. Antennal resolution of pheromone plumes in three moth species. J. Insect Physiol. 48:433-442.

Justus, K.A., J. Murlis, C. Jones and R.T. Cardé. 2002. Measurement of odor-plume structure in a wind tunnel using a photoionization detector and a tracer gas. Environ. Fluid Mech. 2:115-142.

Farrell, J.A., J. Murlis, W. Lei and R.T. Cardé. 2002. Filament-based atmospheric dispersion model to achieve short time-scale structure of odor plumes. Environ. Fluid Mech. 2:143-169.

Li, W. J.A. Farrell and R.T. Cardé. 2001. Tracking of fluid-advected odor plumes: strategies inspired by insect orientation to pheromone. Adapt. Behav. 9:143-167.

Kenna, M.A., W.E. Wallner, P.S. Grinburg and R.T. Cardé. 2001. Female flight propensity in Lymantria dispar (Lepidoptera: Lymantriidae) from Russia, North America and their reciprocal F1 hybrids. Environ. Entomol. 30: 380-387.

Dekker, T., W. Takken and R.T. Cardé. 2001. Structure of host-odour plumes influences catch of Anopheles gambiae s.s. and Aedes aegypti in a dual-choice olfactometer. Physiol. Entomol. 26:124-134.

Murlis, J., M.A. Willis and R.T. Cardé. 2000. Spatial and temporal structures of pheromone plumes in fields and forests. Physiol. Entomol. 25: 211-222.

Cardé, R.T. and B.G.J. Knols. 2000.  Effects of light levels and plume structure on the orientation manoeuvres of male gypsy moths flying along pheromone plumes.  Physiol. Entomol. 25: 141-150.

Edited Books

Allison, J.D. and R.T. Cardé, eds. 2016. Pheromone Communication in Moths. Evolution, Behavior and Application.  University of California Press, Berkeley, 401 p.

Cardé, R.T. and V.H. Resh, eds. 2012. A World of Insects. The Harvard University Press Reader. Harvard University Press. 404 p.

Resh, V.H. and R.T. Cardé, eds. 2009. Encyclopedia of Insects, 2nd edn. Academic Press, San Diego. 1132 p.

Cardé, R.T. and J.G. Millar, eds. 2004. Advances in the Chemical Ecology of Insects. Cambridge University Press, Cambridge. 341 p.

Resh, V.H. and R.T. Cardé, eds. 2003. Encyclopedia of Insects, 1st edn., Academic Press, San Diego. 1266 p.

Cardé, R.T. and A. K. Minks, eds. 1996. Insect Pheromone Research. New Directions. Chapman & Hall, New York. 684 p.

Cardé, R.T. and W.J. Bell, eds. 1995. Chemical Ecology of Insects 2. Chapman & Hall, New York. 433 p.

Bell, W.J. and R.T. Cardé, eds. 1984. Chemical Ecology of Insects. Chapman & Hall, London. 524 p.

Berenbaum, M.R., R.T. Cardé and G.E. Robinson, eds. 1999-2016. Annual Review of Entomology (volumes 44-60).

General Campus Information

University of California, Riverside
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Riverside, CA 92521
Tel: (951) 827-1012

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417 Entomology Bldg.

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