Table of Contents

Prof. Sharoni Shafir Media Gallery

Contact Us

The Department of Entomology
The Robert H. Smith Faculty of Agriculture, Food and Environment
The Hebrew University of Jerusalem
Herzl 229, Rehovot 7610001, ISRAEL

Tel: 08-9489223 
Fax: 08-9366768


Publications in refereed journals:

Please send an email to if you would like a reprint

Hendriksma, H. P. ; Toth, A. L. ; Shafir, S. Individual and Colony Level Foraging Decisions of Bumble Bees and Honey Bees in Relation to Balancing of Nutrient Needs. Frontiers in Ecology and Evolution 2019, 7 177.

Maori, E. ; Garbian, Y. ; Kunik, V. ; Mozes-Koch, R. ; Malka, O. ; Kalev, H. ; Sabath, N. ; Sela, I. ; Shafir, S. A Transmissible RNA Pathway in Honey Bees. Cell Rep 2019, 27, 1949-1959.e6. Abstract

Topman, S., D. Tamir-Ariel, H. Bochnic-Tamir, T. Stern, S. Shafir, S. Burdman, and Z. Hayouka. 2018. Random peptide mixtures as new crop protection agents. Microbial Biotechnology. In press.

Wright, G.A., S.W. Nicolson and S. Shafir. 2018. Nutritional physiology and ecology of honey bees. Annual Review of Entomology 63: 327-344.

Sapir G, Z. Baras, G. Azmon, M. Goldway, S. Shafir, A. Allouche, E. Stern, RA Stern. 2017. Synergistic effects between bumblebees and honey bees in apple orchards increase cross pollination, seed number and fruit size. Scientia Horticulturae 219:107-117. doi: 10.1016/j.scienta.2017.03.010.

Zarchin, S. ; Dag, A. ; Salomon, M. ; Hendriksma, H. P. ; Shafir, S. Honey bees dance faster for pollen that complements colony essential fatty acid deficiency. Behavioral Ecology and Sociobiology 2017, 71, 172.

Hendriksma HP and S. Shafir. 2016. Honey bee foragers balance colony nutritional deficiencies. Behavioral Ecology and Sociobiology 70: 509-517

Arien Y., A. Dag, S. Zarchin, T. Masci and S. Shafir. 2015. Omega-3 deficiency impairs honey bee learning. Proceedings of the National Academy of Sciences of the USA 112: 15761-15766

Nyska R., S. Shafir, M. Goldway and D. Schneider. 2014. Effect of enhanced pollination on economic properties of self-compatible loquat (Eriobotrya japonica Lindl.). Scientia Horticulturae 176: 248-254

Raz, A., R.A. Stern, S. Shafir, M. Goldway. 2014. Reduced yields of 'Earlicot' apricot (Prunus armeniaca) grown in a relatively hot climate and methods to improve flowering intensity and yield. Journal of Horticultural Science and Biotechnology 89: 495-500

Avni, D., H.P. Hendriksma, Z. Uni, A. Dag and S. Shafir. 2014. Nutritional aspects of honey bee-collected pollen and constraints on colony development in the eastern Mediterranean. Journal of Insect Physiology 69: 65-73

Hendriksma H.P., K. Oxman and S. Shafir 2014. Amino acid and carbohydrate tradeoffs by honey bee nectar foragers and their implications for plant-pollinator interactions. Journal of Insect Physiology 69: 56-64

Afik O, Delaplane KS, Shafir S, Moo-Valle H, Quezada-Euan JJ, 2014. Nectar minerals as regulators of flower visitation in stingless bees and nectar hoarding wasps. Journal of Chemical Ecology 40:476-483. doi: 10.1007/s10886-014-0455-8.

Shafir S. and L. Yehonatan. 2013. Comparative evaluations of reward dimensions in honey bees – evidence from 2-alternative forced choice proboscis-extension conditioning. Animal Cognition 17:633-644. doi: 10.1007/s10071-013-0694-z.

Nyska R., A. Raz, Z. Baras, S. Shafir, M. Goldway and D. Schneider. 2013. Self-compatibility in loquat (Eriobotrya japonica Lindl.) is due to S6-RNase mutation. Scientia Horticulturae 161: 43-48.

Fleischer A., S. Shafir, and Y. Mandelik. 2013. A proactive approach for assessing alternative management programs for an invasive alien pollinator species. Ecological Economics 88: 126-132.

Vanbergen A.J., and the Insect Pollinators Initiative: M. Baude, J.C. Biesmeijer, N.F. Britton, M.J.F. Brown, M. Brown, J. Bryden, G.E. Budge, J.C. Bull, C. Carvell, A.J. Challinor, C.N. Connolly, D.J. Evans, E.J. Feil, M.P. Garratt, M.K. Greco, M.S. Heard, V.A.A. Jansen, M.J. Keeling, W.E. Kunin, G.C. Marris, J. Memmott, J.T. Murray, S.W. Nicolson, J.L. Osborne, R.J. Paxton, C.W.W. Pirk, C. Polce, S.G. Potts, N.K. Priest, N.E. Raine, S. Roberts, E.V. Ryabov, S. Shafir, M.D.F. Shirley, S.J. Simpson, P.C. Stevenson, G.N. Stone, M. Termansen, and G.A. Wright. 2013. Threats to an ecosystem service: pressures on pollinators. Frontiers in Ecology and the Environment Frontiers in Ecology and the Environment 11: 251-259.

Garbian, Y., E. Maori, H. Kalev, S. Shafir and I. Sela. 2012. Bidirectional transfer of RNAi between honey bee and Varroa destructor: Varroa gene silencing reduces Varroa population. PLOS Pathogens 8 (12): e1003035.

Moyal Ben Zvi, M., E. Shklarman, T. Masci, H. Kalev, T. Debener, S. Shafir, M. Ovadis and A. Vainstein. 2012. PAP1 transcription factor enhances production of phenylpropanoid and terpenoid scent compounds in rose flowers. New Phytologist 195: 335-345.

Kaspi, R. and S. Shafir. 2012. Associative olfactory learning of the red dwarf honey bee Apis florea. Apidologie 44: 100-109.

Afik, O., A. Dag, Y. Yeselson, A. Schaffer and S. Shafir. 2010. Selection and breeding of honey bees for higher or lower collection of Avocado nectar. Journal of Economic Entomology 103: 228-233.

Shafir, S. and A.B. Barron. 2010. Optic flow informs distance but not profitability for honeybees. Proceedings of the Royal Society Series B 277: 1241-1245.

Afik, O., T. Hallel, A. Dag and S. Shafir. 2009. The components that determine honey bee (Apis mellifera) preference between Israeli unifloral honeys and the implication for nectar attractiveness. Israel Journal of Plant Sciences 57: 253-261.

Avni, D., A. Dag and S. Shafir. 2009. Pollen sources for honey bees in Israel: Source, periods of shortage and influence on population growth. Israel Journal of Plant Sciences 57: 263-275.

Shafir, S., L. Kabanoff, M. Duncan and B. P. Oldroyd. 2009. Honey bee (Apis mellifera) sperm competition in vitro – two are no less viable than one. Apidologie 40: 556-561.

Drezner-Levy, T., B. H. Smith and S. Shafir. 2009. The effect of foraging specialization on various learning tasks in the honey bee (Apis mellifera). Behavioral Ecology and Sociobiology 64: 135-148.

Maori, E., N. Paldi, S. Shafir, H. Kalev, E. Tsur, E. Glick, and I. Sela. 2009. IAPV, a bee-affecting virus associated with colony collapse disorder can be silenced by dsRNA ingestion. Insect Molecular Biology 18: 55-60.

Avni, D., A. Dag, and S. Shafir. 2009. The effect of surface area of pollen patties fed to honey bee (Apis mellifera) colonies on their consumption, brood production and honey yields. Journal of Apicultural Research 48: 23-28.

Sapir, G., R.A. Stern, S. Shafir, and M. Goldway. 2008. S-RNase based S-genotyping of Japanese plum (Prunus salicina Lindl.) and its implication on the assortment of cultivar-couples in the orchard. Scientia Horticulturae 118: 8-13.

Sapir, G., R.A. Stern, S. Shafir, and M. Goldway. 2008. Full compatibility is superior to semi-compatibility for fruit set in Japanese plum (Prunus salicina Lindl.) cultivars. Scientia Horticulturae 116: 394-398.

Shafir S., T. Reich, E. Tsur, I. Erev and A. Lotem. 2008. Perceptual accuracy and conflicting effects of certainty on risk-taking behaviour. Nature 453: 917-920.

Afik, O., A. Dag, and S. Shafir. 2008. Honey bee (Apis mellifera) round dance is influenced by trace elements composition of floral nectar. Animal Behaviour. 75: 371-377.

Sapir, G., R.A. Stern, S. Shafir, and M. Goldway. 2007. SFBs of Japanese plum (Prunus salicina): cloning seven alleles and determining their linkage to the S-RNase gene. HortScience 42: 1509-1512.

Stern, R.A., G. Sapir, S. Shafir, A. Dag, and M. Goldway. 2007. The appropriate management of honey bee colonies for pollination of Rosaceae fruit trees in warm climates. Middle Eastern and Russian Journal of Plant Science and Biotechnology. 1: 13-19.

Sapir, G., M. Goldway, S. Shafir, and R.A. Stern. 2007. Multiple introduction of honey bee colonies increases cross-pollination, fruit-set and yield of ‘Black-

Diamond’ Japanese plum (Prunus salicina L.). Journal of Horticultural Science and Biotechnology. 82: 590-596.

Drezner-Levy, T. and S. Shafir. 2007. Parameters of variable reward distributions that affect risk sensitivity of honey bees. Journal of Experimental Biology. 210: 269-277.

Afik, O., and S. Shafir. 2007. Effect of ambient temperature on crop loading in the honey bee, Apis mellifera (Hymenoptera: Apidae). Entomologia Generalis. 29: 135-148.

Afik, O., A. Dag, and S. Shafir. 2007. Perception of avocado bloom (Lauraceae: Persea americana) by the honey bee (Hymenoptera: Apidae: Apis mellifera). Entomologia Generalis. 30: 135-153.

Shafir, S., A. Dag, A. Bilu, M. Abu-Toamy, and Y. Elad. 2006. Honey bee dispersal of the biocontrol agent Trichoderma harzianum T39: effectiveness in suppressing Botrytis cinerea on strawberry under field conditions. European Journal of Plant Pathology. 116: 119-128.

Afik, O., A. Dag, Z. Kerem, and S. Shafir. 2006. Analyses of avocado (Persea americana) nectar properties and their perception by honey bees (Apis mellifera). Journal of Chemical Ecology. 32: 1949-1963.

Afik, O., A. Dag, and S. Shafir. 2006. The effect of avocado (Persea americana) nectar composition on its attractiveness to honey bees. Apidologie. 37: 317-325.

Dag, A., O. Afik, Yeselson, Y., Schaffer, A., and S. Shafir. 2006. Physical, chemical and palynological characterization of avocado (Persea Americana Mill.) honey in Israel. International Journal of Food Science and Technology. 41: 387-394.

Carter, C., S. Shafir, L. Yehonatan, R. G. Palmer, and R. Thornburg. 2006. A novel role for proline in plant floral nectars. Naturwissenschaften. 93: 72-79.

Zisovich, A. H., R. A. Stern, S. Shafir, and M. Goldway. 2005. Fertilization efficiency of semi- and fully-compatible European pear (Pyrus communis) cultivars. Journal of Horticultural Science and Biotechnology. 80: 143-146.

Dag A., R. A. Stern, S. Shafir. 2005. Honey bee (Apis mellifera) strains differ in apple (Malus domestica) pollen foraging preference. Journal of Apicultural Research 44: 15-20.

Zisovich, A. H., R. A. Stern, G. Sapir, S. Shafir, and M. Goldway. 2004. The RHV region of S-Rnase in the European pear (Pyrus communis) is not required for the determination of specific pollen rejection. Sexual Plant Reproduction 17: 151-156.

Shafir, S., G. Menda and B.H. Smith. 2005. Caste-specific differences in risk-sensitivity in honey bees (Apis mellifera). Animal Behaviour 69: 859-868.

Shalit, M., S. Shafir, O. Larkov, E. Bar, D. Kaslassi, Z. Adam, D. Zamir, A. Vainstein, D. Weiss, U. Ravid and E. Lewinsohn. 2004. Volatile compounds emitted by rose cultivars: Fragrance perception by man and honeybees. Israel Journal of Plant Sciences 52: 245-255.

Stern, R.A., M. Goldway, A. H. Zisovich, S. Shafir and A. Dag. 2004. Sequential introduction of honeybee colonies increases cross-pollination, fruit-set and yield of ‘Spadona’ pear (Pyrus communis L.). Journal of Horticultural Science and Biotechnology 79: 652-658.

Freeman, S., D. Minz, I. Kolesnik, O. Barbul, A. Zveibill, M. Maymon, Y. Nitzani, B. Kirshner, D. Rav-David, A. Bilu, A. Dag, S. Shafir, and Y. Elad. 2004. Trichoderma biocontrol of Colletotrichum acutatum and Botrytis cinerea in strawberry, population survival, and identification of biocontrol isolates according to ITS sequence analysis. European Journal of Plant Pathology 110: 361-370.

Bilu, A., A. Dag, Y. Elad, and S. Shafir. 2004. Honey bee dispersal of biocontrol agents: an evaluation of dispensing devices. Biocontrol Science and Technology 14: 607-617.

Zisovich, A. H., R. A. Stern, S. Shafir, and M. Goldway (PI). 2004. Identification of seven S-alleles from the European pear (Pyrus communis) and the determination of compatibility among cultivars. Journal of Horticultural Science and Biotechnology 79: 101-106.

Weber, E. U., S. Shafir, and A.R. Blais. 2004. Predicting risk sensitivity in humans and lower animals: risk as variance or coefficient of variation. Psychological Review 111: 430-445.

Paldi, N., S. Zilber, and S. Shafir. 2003. Associative olfactory learning of honeybees to differential rewards in proboscis-extension and free-flying behavioral assays. Journal of Chemical Ecology 29: 2515-2538.

London-Shafir, I., S. Shafir, and D. Eisikowitch. 2003. Amygdalin in almond nectar and pollen –facts and possible roles. Plant Systematics and Evolution 238: 87-95.

Shafir, S., A. Bechar, and E. U. Weber. 2003. Cognition-mediated coevolution – context-dependent evaluations and sensitivity of pollinators to variability in nectar rewards. Plant Systematics and Evolution 238: 195-209.

Dag, A., A.E. Fetscher, O. Afik, Y. Yeselson, A.A. Schaffer, Y. Kamer, N.M. Waser, M.A. Madore, M.L. Arpaia, R. Hofshi, and S. Shafir. 2003. Honey bee (Apis mellifera) strains differ in avocado (Persea americana) nectar foraging preferences. Apidologie 34: 299-309.

Kalev, H., A. Dag, and S. Shafir. 2002. Feeding pollen supplements to honey bee colonies during pollination of sweet pepper in enclosures. American Bee Journal 142: 675-679.

Dvash, L, O. Afik, S. Shafir, A. Schaffer, Y. Yeselson, A. Dag, and S. Landau. 2002. Determination by near-infrared spectroscopy of perseitol used as a marker for the botanical origin of Avocado (Persea americana Mill.) honey. Journal of Agricultural and Food Chemistry 50: 5283-5287.

Lavid, N., J. Wang, M. Shalit, I. Guterman, E. Bar, T. Beuerle, N. Menda, S. Shafir, D. Zamir, Z. Adam, A. Vainstein, D. Weiss, E. Pichersky, and E. Levinsohn. 2002. O-Methyltransferases involved in the biosynthesis of volatile phenolic derivatives in rose petals. Plant Physiology. 129: 1899-1907.

Shafir, S., T.A. Waite, and B.H. Smith. 2002. Context-dependent violations of rational choice in honeybees (Apis mellifera) and gray jays. Behavioral Ecology and Sociobiology 51: 180-187.

Shafir, S. 2000. Risk-sensitive foraging: the effect of relative variability. Oikos 89: 663-669.

Shafir, S., D.D. Wiegmann, B.H. Smith, and L.A. Real. 1999. Risk-sensitive foraging: choice behaviour of honeybees in response to variability in volume of reward. Animal Behaviour 57: 1055-1061.

Shafir, S. and J. Roughgarden. 1998. Testing predictions of foraging theory for a sit-and-wait forager, Anolis gingivinus. Behavioral Ecology 9 (1): 74-84.

Shafir, S. 1996. Color discrimination conditioning of a wasp, Polybia occidentalis (Hymenoptera: Vespidae). Biotropica 28 (2): 243-251.

Shafir, S. 1994. Intransitivity of preferences in honey bees -- support for comparative evaluation of foraging options. Animal Behaviour 48: 55-67.

Shafir, S. and J. Roughgarden. 1994. Instrumental discrimination conditioning of Anolis cristatellus in the field with food as a reward. Caribbean Journal of Science 30 (3-4): 228-233.

Wetterer, J., S. Shafir, L. Morrison, K. Lips, G. Gilbert, M. Cipollini, and C. Blaney. 1992. On- and off-trail orientation in the leaf-cutting ant, Atta cephalotes (L.) (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society 65: 96-98.

McClanahan T.R. and S. Shafir. 1990. Causes and consequences of sea urchin abundance and diversity in Kenyan coral reef lagoons. Oecologia 83: 362-370.

Chapters in collections:

Shafir, S. 2011. Bee cognition and crop pollination: proven and potential applications. In: Seckbach J. and Z. Dubinsky, eds., All flesh is grass: plant-animal interactions. pp. 185-198.

Shafir, S. and A. Dag. 2008. Honey. In: Reifen R., and G. Rozen, eds., The intelligence of nutrition (In Hebrew). pp. 392-395.

Shafir, S. and O. Afik. 2000. The effect of ambient temperature on crop load size in honey bees. In: Sommeijer, M. and A. de Ruijter, eds., Insect Pollination in Greenhouses, CIP-DATA Koninklijke Bibliotheek, Den Haag. pp. 155-164.

Shafir, S. and J. Roughgarden. 1996. The effect of memory length on individual fitness in a lizard. In R.K. Belew and M. Mitchell, eds., Adapting Individuals in Evolving Populations: Models and Algorithms, (Santa Fe Institute Studies in the Sciences of Complexity), Addison Wesley, Reading, MA. pp. 173-182.

Shafir, S. 1996. Preface to Peter Todd's “Sexual selection and the evolution of learning”. In R.K. Belew and M. Mitchell, eds., Adapting Individuals in Evolving Populations: Models and Algorithms, (Santa Fe Institute Studies in the Sciences of Complexity), Addison Wesley, Reading, MA. pp. 361-364.

Roughgarden, J., A. Bergman, S. Shafir and C. Taylor. 1996. Adaptive Computation in Ecology and Evolution: A Guide for Future Research. In R.K. Belew and M. Mitchell, eds., Adapting Individuals in Evolving Populations: Models and Algorithms, (Santa Fe Institute Studies in the Sciences of Complexity), Addison Wesley, Reading, MA. pp. 25-30.

Shafir, S. and J. Roughgarden . 1994. The effect of memory length on the foraging behavior of a lizard. In D. Cliff, P. Husbands, J-A Meyer and S.W. Wilson, eds., From Animals to Animats 3 – Proceedings of the Third International Conference on Simulation of Adaptive Behavior, MIT Press, Cambridge, MA, pp. 221-225.