DANIEL RICHARD PAPAJ
Professor and Associate Head
Department of Ecology & Evolutionary Biology
University of Arizona
Tucson, AZ 85721 U.S.A.
PHONE NUMBER:
(520) 621-8988 (office)
(520) 621-9190 (FAX)
E-MAIL ADDRESS:
papaj@email.arizona.edu
CHRONOLOGY OF EDUCATION
Cornell University, Ithaca, New York
B.S. in Biology with honors, received June 1978.
Duke University, Durham, North Carolina
Ph.D. in Zoology, received November 1984.
Dissertation Title: Causes of variation in host discrimination in the butterfly, Battus philenor.
Major Advisor: Mark D. Rausher.
Major Fields: Animal Behavior, Insect Behavior, Behavioral Ecology, Insect-Host Interactions,
Thermal Ecology
PUBLICATIONS
Books and Monographs
D.R. Papaj and A.C. Lewis (eds). 1993. Insect Learning: Ecological and Evolutionary
Perspectives. Chapman and Hall, Inc. New York.
Book Chapters
J. Jaenike and D.R. Papaj. 1992. Learning and patterns of host use by insects. pp. 245-264 In
Chemical Ecology: An Evolutionary Perspective. M.B. Isman and B.D. Roitberg, eds. Chapman
and Hall, Inc., New York.
D.R. Papaj. 1993. Use and avoidance of occupied hosts as a dynamic process in tephritid flies. In
Insect-Plant Interactions. Vol. 5. E.A. Bernays, ed. CRC Press. Boca Raton, Florida.
D.R. Papaj. 1993. The evolution of instinct: lessons from learning in parasitoids. In Insect
Learning: Ecological and Evolutionary Perspectives. Chapman and Hall, Inc., D.R. Papaj and
A.C. Lewis, eds. Chapman and Hall, Inc. New York.
2/12/2016
D.R. Papaj/2
D.R. Papaj 1994. Optimizing learning and its effects on evolutionary change. pp. 133-153 in
Behavioral Mechanisms in Evolutionary Ecology. L. Real, ed. University of Chicago Press.
Chicago, Illinois.
D.R. Papaj. 1997. How a butterfly copes with the problem of biodiversity. In Ecological
Perspectives of Biodiversity, T. Abe, M. Higashi and S. Levin, eds. Springer-Verlag, Berlin.
R.J. Prokopy and D.R. Papaj. 1999. Behavior of flies of the genera Rhagoletis, Zonosemata and
Carpomya (Trypetinae: Carpomyina). pp. 219-252 in Fruit Flies (Tephritidae): Phylogeny and
Evolution of Behavior. M. Aluja and A.L. Norrbom (eds.). CRC Press. Boca Raton, Florida.
944p.
F. Diaz-Fleischer, D.R. Papaj, R.J. Prokopy, A.L. Norrbom and M. Aluja. 1999. Evolution of
fruit fly oviposition behavior. pp. 811-842 in Fruit Flies (Tephritidae): Phylogeny and
Evolution of Behavior. M. Aluja and A.L. Norrbom (eds.). CRC Press. Boca Raton, Florida.
944p.
Reviews
D.R. Papaj and R.J. Prokopy. 1989. Ecological and evolutionary aspects of learning in
phytophagous insects. Annual Review of Entomology 34:315-350.
D.R. Papaj. 2000. Ovarian dynamics and host use. Annual Review of Entomology 45:423-448.
C. Nufio and D.R. Papaj. 2001. Host-marking behavior in phytophagous insects and parasitoids.
Entomologia Experimentalis et Applicata 99:273-293.
D.R. Papaj. 2003. Learning. In Encyclopedia of Insect Learning, V. Resh and R. Carde, eds.
Academic Press. New York.
Book Reviews
D.R. Papaj. 1993. The Tinbergen Legacy. M.S. Dawkins, T.R. Halliday and R. Dawkins, eds.
American Scientist 81:390-391.
D.R. Papaj. 1998. Comparative Psychology of Invertebrates: The Field and Laboratory Study of
Insect Behavior. G. Greenberg and E. Tobach, eds. Quarterly Review of Biology, 73:250.
Refereed Journal Articles
D. Pimentel, D. Nafus, W. Vergara, D. Papaj, L. Jaconetta, M. Wulfe, L. Olsvig, M. Loye and E.
Mendoza. 1978. Biological solar energy conversion and U.S. energy policy. Bioscience 28:376382.
M.D. Rausher and D.R. Papaj. 1983. Foraging by Battus philenor butterflies: evidence for
individual differences in searching behaviour. Animal Behaviour 31:341-347.
M.D. Rausher and D.R. Papaj. 1983. Demographic consequences of conspecific host
discrmination by Battus philenor butterflies. Ecology 64:1402-1410.
2/12/2016
D.R. Papaj/3
R.J. Prokopy, D.R. Papaj, S.S. Cooley, and C. Kallet. 1985. On the nature of learning in
oviposition site acceptance by apple maggot flies. Animal Behaviour 34:98-107.
R.J. Prokopy, D.R. Papaj and T.T.Y. Wong. 1986. Fruit foraging behavior of Mediterranean fruit
fly females on host and non-host plants. Florida Entomologist 69:651-657.
D.R. Papaj. 1986. Shifts in foraging behavior by a Battus philenor population: field evidence for
switching by individual butterflies. Behavioral Ecology and Sociobiology 19:31-39.
D.R. Papaj. 1986. Leaf buds: a factor in host selection by Battus philenor butterflies. Ecological
Entomology 11:301-307.
D.R. Papaj. 1986. Conditioning of leaf-shape discrimination by chemical cues in the butterfly,
Battus philenor. Animal Behaviour 34:1281-1288.
D.R. Papaj. 1986. Inter-population differences in host preference and the evolution of learning in
the butterfly, Battus philenor. Evolution 40:518-530.
D.R. Papaj and R.J. Prokopy. 1986. The phytochemical basis of learning in Rhagoletis pomonella
and other herbivorous insects. Journal of Chemical Ecology 12:1125-1143.
R.J. Prokopy, D.R. Papaj, S.B. Opp and T.T.Y. Wong. 1987. Intra-tree foraging behavior of
Ceratitis capitata flies in relation to host fruit density and quality. Entomologia Experimentalis et
Applicata 45:251-258.
D.R. Papaj and M.D. Rausher. 1987. Components of conspecific host plant discrimination by
Battus philenor (Papilionidae). Ecology 68:245-253.
D.R. Papaj and M.D. Rausher. 1987. Genetic differences and phenotypic plasticity as causes of
variation in oviposition preference in Battus philenor. Oecologia 74:24-30.
D.R. Papaj, R.J. Prokopy, P. McDonald and T.T.Y. Wong. 1987. Differences in learning between
laboratory and wild Ceratitis capitata flies. Entomologia Experimentalis et Applicata 45:65-72.
D.R. Papaj and R.J. Prokopy. 1988. The effect of prior adult experience on components of
habitat preference in the apple maggot fly (Rhagoletis pomonella). Oecologia 76:538-543.
R.J. Prokopy and D.R. Papaj. 1988. Learning of apple fruit biotypes by apple maggot flies.
Journal of Insect Behavior 1:67-74.
R.J. Prokopy, M. Aluja, D.R. Papaj, B.D. Roitberg and T.T.Y. Wong. 1989. Influence of
previous experience with host plant foliage on behavior of Mediterranean fruit fly females.
Proceedings of the Hawaiian Entomological Society 29:97-101.
D.R. Papaj, S.B. Opp, R.J. Prokopy and T.T.Y. Wong. 1989. Cross-induction of host fruit
acceptance in medfly: the role of fruit size and chemistry. Journal of Insect Behavior 2:341-354.
2/12/2016
D.R. Papaj/4
D.R. Papaj, J. Hendrichs and B.I. Katsoyannos. 1989. Use of fruit wounds in oviposition by
Mediterranean fruit flies. Entomologia Experimentalis et Applicata 53:203-209.
D.R. Papaj, B.D. Roitberg and S.B. Opp. 1989. Serial effects of host infestation on egg allocation
by the Mediterranean fruit fly: a rule of thumb and its functional significance. Journal of Animal
Ecology 58:955-970.
D.R. Papaj. 1990. Fruit size and clutch size in Ceratitis capitata. Entomologia Experimentalis et
Applicata 54:195-198.
M.R. Strand, B.D. Roitberg and D.R. Papaj. 1990. Acridine orange: a potentially useful internal
marker of Hymenoptera and Diptera. Journal of the Kansas Entomological Society 63:634-637.
W.J. Lewis, L.E.M. Vet, J.H. Tumlinson, J.C. van Lenteren and D.R. Papaj. 1990. Variation in
parasitoid foraging behavior: essential element of a sound biological control theory.
Environmental Entomology 19:1183-1193.
L.E.M. Vet, W.J. Lewis, D.R. Papaj., and J.C. van Lenteren. 1990. A variable-response model
for parasitoid foraging behaviour. Journal of Insect Behavior 3:471-490.
D.R. Papaj, B.D. Roitberg, S.B. Opp, M. Aluja, R.J. Prokopy and T.T.Y. Wong. 1990. Effect of
marking pheromone on clutch size in the Mediterranean fruit fly. Physiological Entomology
15:463-468.
D.R. Papaj and L.E.M. Vet. 1990. Odor learning and foraging success in the parasitoid,
Leptopilina heterotoma. Journal of Chemical Ecology 16:3137-3150.
J. Hendrichs, B.I. Katsoyannos, D.R. Papaj and R.J. Prokopy. 1991. Mediterranean fruit fly
(Diptera: Tephritidae) activities in nature, including feeding and fecundity on natural food
sources. Oecologia 86:223-231.
D.R. Papaj, P. Feeny, K. Sachdev and L. Rosenberry. 1992. D-(+)-pinitol, an oviposition
stimulant for the pipevine swallowtail butterfly (Battus philenor). Journal of Chemical Ecology
18:799-815.
L.E.M. Vet and D.R. Papaj. 1992. Effects of experience on parasitoid movement in odour
plumes. Physiological Entomology 17:90-96.
R.J. Prokopy, D.R. Papaj, J. Hendrichs and T.T.Y. Wong. 1992. Behavioral responses of
Ceratitis capitata flies to bait spray droplets and natural food. Entomologia Experimentalis et
Applicata 64:247-257.
D.R. Papaj, A.L. Averill, R.J. Prokopy and T.T.Y. Wong. 1992. Host-marking pheromone and
use of previously-established oviposition sites by the Mediterranean fruit fly. Journal of Insect
Behavior 5:583-598.
2/12/2016
D.R. Papaj/5
R.J. Prokopy, S.S. Cooley and D.R. Papaj. 1993. How well can relative specialist Rhagoletis flies
learn to discriminate fruit for oviposition? Journal of Insect Behavior 6:167-176.
D.R. Papaj and M. Aluja. 1993. Temporal dynamics of host-marking in the tropical tephritid fly,
Anastrepha ludens. Physiological Entomology 18:279-284.
D.R. Papaj, H. Snellen, K. Swaans and L.E.M. Vet. 1994. Unrewarding experiences and their
effect on foraging in parasitic wasps. Journal of Insect Behavior 7:465-481.
Papaj, D.R. 1994. Oviposition site guarding by male walnut flies and its possible consequences
for mating success. Behavioral Ecology and Sociobiology 34:187-195.
M.L. Henneman, D.R. Papaj, A.J. Figueredo and L.E.M. Vet. 1995. Egg-laying experience and
acceptance of parasitized hosts by the parasitoid, Leptopilina heterotoma (Hymenoptera:
Eucoilidae). Journal of Insect Behavior 8:331-342.
Papaj, D.R., J.M. García and H. Alonso-Pimentel. 1996. Marking of host fruit by male
Rhagoletis boycei Cresson flies (Diptera: Tephritidae) and its effect on egg-laying. Journal of
Insect Behavior 9:585-598.
D.R. Papaj and R.H. Messing. 1996. Functional shifts in the use of parasitized hosts by a
tephritid fly: The role of host quality. Behavioral Ecology 7:235-242.
H. Alonso-Pimentel and D.R. Papaj. 1996. Operational sex ratio vs. gender densities as
determinants of copulation duration in the walnut fly, Rhagoletis juglandis (Diptera:
Tephritidae). Behavioral Ecology and Sociobiology 39:171-180.
H. Alonso-Pimentel and D.R. Papaj. 1996. Patterns of egg load and their significance for the
distribution of a tephritid fly in time and space. Annals of the Entomological Society of America
89:875-882.
D.R. Papaj and H. Alonso-Pimentel. 1997. Why walnut flies superparasitize: time savings as a
possible function. Oecologia 109:166-174.
R.A. Allard and D.R. Papaj. 1997. Learning of visual cues by pipevine swallowtail butterflies: A
test using artificial leaf models. Journal of Insect Behavior 9:961-967.
L.E.M. Vet, A.G. De Jong, E. Franchi, and D.R. Papaj. 1998. The effect of complete versus
incomplete information on odour discrimination in a parasitic wasp. Animal Behaviour 55:12711279.
D.R. Papaj and R.H. Messing. 1998. Asymmetries in dynamical state as an explanation for
resident advantage in contests. Behaviour 135:1013-1030.
H. Alonso-Pimentel, J.B. Korer, C. Nufio and D.R. Papaj. 1998. Role of color and shape stimuli
in host-enhanced oogenesis in the walnut fly, Rhagoletis juglandis. Physiological Entomology
23:97-104.
2/12/2016
D.R. Papaj/6
H. Alonso-Pimentel and D.R. Papaj. 1999. The interaction between effects of resource presence
and sex ratio on copulation duration in a walnut fly, Rhagoletis juglandis (Diptera: Tephritidae).
Animal Behavior 57:1063-1069.
M.L. Henneman and D.R. Papaj. 1999. Role of host fruit color pattern in the behavior of
Rhagoletis juglandis (Diptera: Tephritidae). Entomologia Experimentalis et Applicata 93:249258.
H. Alonso-Pimentel, H.G. Spangler, R. Rogers and D.R. Papaj. 2000. Acoustic component and
social context of the wing display of the walnut fly Rhagoletis juglandis. Journal of Insect
Behavior 13:511-524.
C.R. Nufio, D. R. Papaj and H. Alonso-Pimentel. 2000. A field analysis of host fruit utilization
by the walnut fly, Rhagoletis juglandis. Environmental Entomology 93:994-1001.
A.D. Lachmann and D.R. Papaj. 2001. Effect of host stimuli on ovariole development in the
walnut fly, Rhagoletis juglandis (Diptera, Tephritidae). Physiological Entomology 26:38-48.
M. Aluja, F. Díaz-Fleischer, D.R. Papaj, G. Lagunes and J. Sivinski. 2001. Effects of age, diet,
female density, and the host resource on egg load in Anastrepha ludens and A. obliqua (Diptera:
Tephritidae). Journal of Insect Physiology 47:975-988.
R. Dukas, R.J. Prokopy, D.R. Papaj and J.J. Duan. 2001. Egg-laying behavior of Mediterranean
fruit flies: is social facilitation important? Florida Entomologist 84:665-671.
M.R. Weiss and D.R. Papaj. 2003. Butterfly color learning in two different behavioral contexts:
How much can a butterfly keep in mind? Animal Behaviour 65: 425-434.
J. Cnaani, J.O. Schmidt and D.R. Papaj. 2003. The effect of octopamine on behavioral responses
of free-foraging bumblebees to a change in food source profitability. Naturwissenschaften
90:185-188.
C.R. Nufio and D.R. Papaj. 2004. Host marking behaviour as a quantitative signal of competition
in the walnut fly Rhagoletis juglandis. Ecological Entomology 29: 336-344.
C.R. Nufio and D.R. Papaj. 2004. Superparasitism of larval hosts by the walnut fly, Rhagoletis
juglandis, and its implications for female and offspring performance. Oecologia 141: 460-467.
D.R. Papaj. 2004. In praise of a good colleague: Ronald John Prokopy. Journal of Insect
Behavior 17: 569-577.
E.A. Hebets and D.R. Papaj. 2005. Complex signal function: Developing a framework for
testable hypotheses. Behavioral Ecology and Sociobiology 57:197-214.
2/12/2016
D.R. Papaj/7
L.D. Carsten and D.R. Papaj. 2005. Effects of reproductive state and host resource experience on
mating decisions in a walnut-infesting fly (Rhagoletis juglandis). Behavioral Ecology 16:528–
533.
B.D. Worden, A.K. Skemp and D.R. Papaj. 2005. Learning in two contexts: the effects of
interference and body size in bumblebees. Journal of Experimental Biology 208:2045-2053.
S.K. Lynn, J. Cnaani, and D.R. Papaj. 2005. Peak shift discrimination learning as a mechanism
of signal evolution. Evolution 59:1300–1305. doi: 10.1554/04-284
D.R. Papaj. 2005. Ovarian dynamics in relation to host quality in the walnut-infesting fly,
Rhagoletis juglandis. Functional Ecology 19:396-404.
D.R. Papaj and G.M. Newsom. 2005. A within-species warning function for an aposematic
signal. Proceedings of the Royal Society of London, Series B 272:2519–2523. doi:
10.1098/rspb.2005.3186
B.D. Worden and D.R. Papaj. 2005. Flower choice copying in bumblebees. Biology Letters 1(4):
504-507, doi: 10.1098/rsbl.2005.0368.
J. Cnaani, J.D. Thomson and D.R. Papaj. 2006. The effect of reward properties on learning and
choice in foraging bumblebees. Ethology 112: 278–285. doi: 10.1111/j.1439-0310.2005.01174.x
E.C. Snell-Rood and D.R. Papaj. 2006. Learning signals within sensory environments: Does host
cue learning in butterflies depend on background? Animal Biology 56:173-192.
K.L. Prudic, A.K. Skemp, and D.R. Papaj. 2007. Aposematic coloration, luminance contrast,
and the benefits of conspicuousness. Behavioral Ecology 18:41-46.
D.R. Papaj, H.S. Mallory, and C.A. Heinz. 2007. Extreme weather change and host selection in
the pipevine swallowtail, Battus philenor. Oecologia 152:365-375. doi: 10.1007/s00442-0070658-6
I. Kulahci, A. Dornhaus and D.R. Papaj. Multimodal signals enhance decision making in
foraging bumblebees. Proceedings of the Royal Society of London, in press.
E.C. Snell-Rood and D.R. Papaj. 2009. Patterns of phenotypic plasticity in common and rare
environments: A study of host use and color learning in the cabbage white butterfly, Pieris
rapae. The American Naturalist 173: 615–631. doi: 10.1086/597609
E.C. Snell-Rood, W. Gronenberg and D.R. Papaj. 2009. Brain size: A global or induced cost of
learning? Brain, Behavior and Evolution 73:111–128. doi: 10.1159/000213647
2/12/2016
D.R. Papaj/8
WORK SUBMITTED OR IN PREPARATION.
L. Carsten and D.R. Papaj. Individual adjustment of testes size in response to operational sex
ratio. Manuscript completed. Behavioral Ecology, in review.
L. Carsten, D. O’Brien and D.R. Papaj. Adult versus larval orgin of carbon and nitrogen in the
testes of walnut flies. Functional Ecology, in review.
2/12/2016