Prof. David Wool

מחלקה לזואולוגיה אמריטוס
Prof. David Wool
Phone: 03-6409807
Fax: 03-6409403
Office: Sherman - Life Sciences, 310


Period (dates) Name of University Subject Degree or award
1961-1964 Tel Aviv University Biology B.Sc.
1964-1966 Tel Aviv University Zoology M.Sc. (Hons.)

University of Kansas,

Lawrence, USA

Entomology Ph.D.

Title of Master's dissertation: Chironomidae (Diptera) from the Hula Nature Reserve, Israel.
Supervisor: Prof. J. Kugler

Title of Doctoral dissertation: Deviations of zygotic frequencies from expectations in eggs ofTribolium castaneum
Supervisor: Prof. R.R. Sokal




Period (dates) Name of Institution Department Rank / Function
1970-1974 Tel Aviv University Zoology Lecturer
1974-1976 Tel Aviv University Zoology Senior Lecturer

University of Reading,


Agriculture Sabbatical
1977-1981 Tel Aviv University Zoology Senior Lecturer
1981-l988 Tel Aviv University Zoology Associate Professor

Michigan State Univ.,


Zoology Sabbatical
1988-2001 Tel Aviv University Zoology Professor

Macquarie University,

Sydney, Australia

Biol. Sci. Sabbatical
2001-present Tel Aviv University Zoology Professor Emeritus
l990-1992 Tel Aviv University Zoology Department Chairman




  1982-1985 BARD ($150,000)
   Genetic manipulation of almond-moth populations as a means of reducing or preventing insecticide resistance

  1986-l990 CDR-AID ($147,000)
   Identification and characterization of genetic strains in Bemisia tabaci




  •   Entomological Society of America
  •   Society for Population Ecology (Japan)
  •   Israel Zoological Society
  •   Israel Entomological Society
  •   Israel Ecological Society

Research Interests

Ecological and population genetics; evolution;
Biology and ecology of gall-forming aphids.

Recent Publications

Wool, D. The driving forces in Evolution - genetical processes in populations. Ramot Publ. Co. l985 (in Hebrew).


Wool, D. The Driving Forces in Evolution – genetical processes in populations Science Publishers, Enfield, NH , 2006


Refereed Articles

Kugler, J. and D. Wool. Chironomidae (Diptera) from the Hula Nature Preserve, Israel. Ann. Zool. Fenn. 5: 76-83, 1968.

Wool, D. and J. Kugler. Circadian rhythm in Chironomid species (Diptera) from the Hula Nature Preserve, Israel. Ann. Zool. Fenn. 6: 94-97, 1969.


Wool, D. The effect of larval age range on survival of two Tribolium castaneum strains in pure and mixed cultures. Res. Popul. Ecol. 11: 40-44, 1969.

Wool, D. Depth distribution of adults and immatures of two Tribolium castaneum strains in pure and mixed cultures. Res. Popul. Ecol. 11: 137-149, 1969.

Wool, D. Differences in population parameters of two Tribolium castaneum strains in environments of different shapes. Res. Popul. Ecol. 11: 45-66, 1969.

Wool, D. Deviation of zygotic frequencies from expectation in eggs of Tribolium castaneum. Genetics 66: 115--132, 1970.

Sokal, R.R., E.H. Bryant and D. Wool. Selection for changes in genetic facilitation: negative results in Tribolium and Musca. Heredity 25: 299-306, 1970.


Wool, D. A new approach to insecticide resistance: Ecological genetic speculation. Bull. Ent. Soc. Amer. 17: 133-135, 1971.

Wool, D. A critical examination of the use of the Jolicoeur-Brunel diagrams to demonstrate selection in biological populations. Genetics 70: 181-185, 1972.

Wool, D. The growth phase of serially-transferred Tribolium populations. J. Anim. Ecol. 41: 439-451, 1972.

Wool, D. Size, productivity, age and competition in Tribolium populations subjected to long-term serial transfer. J. Anim. Ecol. 42: 183-200, 1973.

Wool, D. Quantitative evaluation of temporal similarity in abundance of animal species. Res. Popul. Ecol. 15: 90-98, 1973.

Sverdlov, E. and D. Wool. The effect of rearing Tribolium castaneum in used culture medium on developmental time of the next generation. Entomol. exp. Appl. 6: 550-551, 1973.

Wool, D. and S. Mendlinger. The eu mutant of the flour beetle, Tribolium castaneum. Environmental and genetic effects on penetrance. Genetica 44: 496-504, 1973.

Wool, D. On the possibility of controlling disease vectors by genetic means. Ha'Refua 84: 658-662, 1973. (in Hebrew).

Sverdlov, E. and D. Wool. Some aspects of survival of starved adult Tribolium castaneum (Herbst). J. Stored Prod. Res. 11: 145-154, 1975.

Wool, D. and E. Sverdlov. Sib-mating populations in an unpredicatable environment: effects on components of fitness. Evolution 30: 119-129, 1976.

Wool, D. Changes of life-history stage distribution of single-strain and mixed Tribolium populations during a single generation at a lowered temperature (Coleoptera, Tenebrionidae). J. Anim. Ecol. 45: 381-394, 1975.

Cohen, E., E. Sverdlov and D. Wool. Expression of Esterases during ontogenesis of the flour beetle, Tribolium castaneum (Tenebrionidae, Coleoptera). Biochem. Genet. 15: 253-264, 1977.

Wool, D. Genetic and environmental components of morphological variation in gall-forming aphids (Homoptera, Aphididae, Fordinae) in relation to climate. J. Anim. Ecol. 46: 875-889, 1977.

Koach, J. and D. Wool. Geographic distribution and host specificity of gall forming aphids (Homoptera, Fordinae) on Pistacia trees in Israel.
Marcellia 40: 207-216, 1977.


Wool, D., H.F. van Emden and S.W. Bunting. Electrophoretic detection of the internal parasite, Aphidius matricariae, in Myzus persicae. Ann. Appl. Biol. 90: 21-26, 1978.

Wool, D., S. Bunting and H.F. van Emden. Electrophoretic study of genetic variation in British Myzus persicae (Sulz.) (Hemiptera, Aphididae). Biochem. Genet. 16: 987-1006, 1978.

Wool, D. and E. Sverdlov. Temporal patterns of food availability and their effect on Tribolium populations. Res. Popul. Ecol. 20: 61-73, 1978.


Wool, D. and O. Bergerson. Analysis of selection processes using the incompletely-penetrant mutant eu of Tribolium castaneum. Can. J. Genet. Cytol. 21: 405-415, 1979.

Wool, D. and O. Bergerson. Sperm precedence in repeated mating of Tribolium adults. (Coleoptera, Tenebrionidae).
Entomol. exp. Appl. 26: 157-160, 1979.

Wool, D. On ecological inference from kurtosis and skewness of morphological characters. Res. Popul. Ecol. 22: 263-272, 1980.

Wool, D. and S. Noiman. Amylase activity in adults and immatures of Tribolium confusum (Coleoptera, Tenebrionidae). Z. Ang. Ent. 90: 382-390, 1980.

Wool, D. and O. Manheim. Genetically-induced susceptibility to malathion in Tribolium castaneum despite selection for resistance. Entomol. exp. Appl. 28: 183-190, 1980.

Nevo, E., T. Perl, A. Beiles, D. Wool & U. Zoller. Genetic structure as a potential monitor of marine pollution. Journees Etud. Pollutions, Cagliari, C.I.S.E.M. pp. 61-68, 1980.

Wool, D. and S. Mendlinger. Genetical and ecological consequences of subdivision in Tribolium populations. J. Anim. Ecol. 50: 421-433, 1981.

Wool, D. and H.F. van Emden. A possible genetic component in the adaptibility of Myzus persicae clones to artificial diet. Z. Ang. Ent. 91: 225-231, 1981.

Nevo, E., T. Perl, A. Beiles and D. Wool. Mercury selection of allozyme genotypes in shrimps. Experientia 37: 1152-1154, 1981.

Wool, D. Critical examination of postulated cladistic relationships among species of flour beetles (Gen. Tribolium; Tenebrionidae, Coleoptera). Biochem. Genet. 20: 333-349, 1982.

Wool, D. Family selection for DDT resistance in flour beetles (Tribolium; Tenebrionidae) as a model for group selection of fitness traits. Isr. J. Entomol. 16: 73-85, 1982.

Noiman, S. and D. Wool. Genetic and ecological properties of malathion resistance in a "field" strain of the flour beetle, Tribolium castaneum (Coleoptera, Tenebrionidae). Z. Ang. Ent. 93: 496-503, 1982.

Graur, D. and D. Wool. Dynamics and genetics of mating behavior in Tribolium (Coleoptera, Tenebrionidae). Behav. Genet. 12: 161-179, 1982.

Wool, D., S. Noiman, O. Manheim and E. Cohen. Malathion resistance in Tribolium strains and their hybrids: Inheritance patterns and possible enzymatic mechanisms. Biochem. Genet. 20: 621-636, 1982.

Wool, D. and S. Noiman. Integrated control of insecticide resistance by combined genetic and chemical treatments: a warehouse model with flour beetles (Tribolium; Tenebrionidae, Coleoptera). Z. Ang. Ent. 95: 22-29, 1983.

Wool, D. and N. Kamin-Belsky. Age-dependent resistance to malathion in adult almond moths, Ephestia cautella (Walker). Z. Ang. Ent. 96: 386-391, 1983.

Wool, D. and O. Manheim. The effects of environment subdivision on morphological variation in the "cauliflower" galls of the aphid Slavum wertheimae (Homoptera, Aphididae, Fordinae). Isr. J. Entomol. 17: 95-1O4, 1983.

Wool, D. Directional and correlated effects of selection on amylase activity, weight and developmental time in Tribolium confusum (Coleoptera, Tenebrionidae). Genetica 65: 173-178, 1984.

Wool, D. and N. Kamin-Belsky. Effects of diet and of larval density on adult sensitivity to malathion and on ecological parameters in Ephestia cautella (Walker) (Lepidoptera, Phyticidae). Z. Ang. Ent. 98: 58-62, 1984.

Wool, D. and E. Shirtz. Selection for high and low amylase activity in adult flour beetles (Tribolium confusum). (Coleoptera, Tenebrionidae). Genetica 63: 229-236, 1984.


Wool, D., D. Gerling and Isabelle Cohen. Electrophoretic detection of two endoparasite species, Encarsia lutea and Eretmocerus mundus, in the whitefly Bemisia tabaci. Z. Ang. Ent. 98: 276-279, 1984.

Wool, D. Blind flour beetles: the expression of variable eye phenotypes in the microcephalic (mc) mutant of Tribolium castaneum (Coleoptera, Tenebrionidae). Isr. J. Entomol. 19: 201-210, 1985.

Graur, D. and D. Wool. Unidirectional interspecific mating in Tribolium castaneum and T. confusum: Evolutionary and ecological implications. Entomol. exp. Appl. 38: 261-265, 1985.

Wool, D. Enzyme activity levels in individuals: Selective value, over-reaction and conditional neutrality.
Evolutionary Theory 8: 39-48, 1986.

Wool, D. and O. Manheim. Population ecology of the gall-forming aphid, Aploneura lentisci in Israel. Res. Popul. Ecol. 28: 151-162, 1986.


Wool, D., Z. Namir and O. Bergerson. Dietary regulation of amylase activity levels in flour beetles (Tribolium. Coleoptera, Tenebrionidae). Ann. Ent. Soc. Amer. 79: 407-413, 1986.

Bergerson, O. and D. Wool. Genetic variation and the ability to colonize new niches in the flour beetle Tribolium castaneum (Herbst) (Coleoptera, Tenebrionidae). Heredity 57: 403-406, 1986.

Wool, D.and O. Bergerson. Random environmental variation and inbreeding: effect on pure-strain and hybrid populations of flour beetles (Tribolium). Can. J. Genet. Cytol. 28: 889-898, 1986.

Wool, D. Differentiation of island populations: a laboratory model. Am. Nat. 129: 188-202, 1987.

Bergerson, O. and D. Wool. Attraction of flour beetles (Tribolium) to wheat flour: heritable character or "conditioning"? Z. Ang. Ent. 104: 179-186, 1987.


Wool, D., J.H. Brower and N. Kamin-Belsky. The relative importance of factors affecting the size of laboratory populations of the almond moth Ephestia cautella (Walker) (Lepidoptera, Pyralidae). Z. Ang. Ent. 104: 217-227, 1987.

Wool, D. and O. Manheim. The effect of host plant properties on gall density, gall weight and clone size in the aphid, Aploneura lentisci (Pass.) (Aphididae, Fordinae) in Israel. Res. Popul. Ecol. 30: 227-234, 1988.

Bergerson, O. and D. Wool. The process of adaptation of flour beetles to new environments. Genetica 77: 3-13, 1988.

Wool, D., J.H. Brower and N. Kamin-Belsky.  The genetic impact of immigrant males on resident populations of the Almond moth, Ephestia cautella (Walker) (Lep., Pyralidae). J. Appl. Ent. 106: 339-344, 1988.

Aloni, R., D. Katz and D. Wool. Effect of the gall-forming aphid Slavum wertheimae on the differentiation of xylem in branches of Pistacia atlantica. Ann. Bot. 63: 373-375, 1989.

Wool, D., D. Gerling, B. Nolt, L.M. Constantino, A. Bellotti and F. Morales. The use of electrophoresis for identification of adult whiteflies (Aleyrodidae) in Israel and Colombia. J. Appl. Ent. 107: 344-350, 1989.

Wool, D. Regular alternation of high and low population size of gall-forming aphids: analysis of ten years of data. Oikos 57: 73-79, 1990.

Wool, D. and S. Greenberg. Esterase activity in whiteflies (Bemisia tabaci) in Israel in relation to insecticide resistance.
Entomol. exp. Appl. 57: 251-258, 1990.


Kamin-Belsky, N. and D. Wool. Dietary modification of digestive physiology in larvae of the almond moth, Ephestia cautella (Walker) (Lepidoptera, Phyticidae). J. Econ. Entomol. 84: 768-775, 1991.

Wool, D. and M. Burstein. A galling aphid with extra life-cycle complexity: population ecology and evolutionary considerations.
Res. Popul. Ecol. 33: 307,322, 1991.


Wool, D. and M. Burstein. Parasitoids attacking the gall-forming aphid, Smynthurodes betae Westw. (Aphidoidea: Fordinae) in Israel. Entomophaga 36: 531-538, 1991.

Wool, D., D. Gerling, A.C. Bellotti, F.J. Morales and B. Nolt. Spatial and temporal genetic variation in populations of the whitefly Bemisia tabaci (Genn.) in Israel and Colombia: an interim report. Insect. Sci. Appl. 12: 225-230, 1991.

Wool, D. and N. Kamin-Belsky. Relative fitness in an ecological context: a three-dimensional representation of fitness of almond moth strains Ephestia cautella (Walker), Lepidoptera, Phyticidae). J. Appl. Ent. 113: 168-174, l992.

Kaufman, B. and D. Wool. Gene flow by immigrants into isolated recipient populations: a laboratory model using flour beetles. Genetica, 85: 163-171, 1992.


Burstein, M. and D. Wool. Great Tits exploit aphid galls as a source of food. Ornis, 23: 107-109, 1992.

Wool, D., J.H. Brower and N. Kamin-Belsky. Reduction of Malathion resistance in caged almond moth Cadra cautella (Walker) (Lepidoptera, Pyralidae) populations by the introduction of susceptible males. J. Stored Prod. Res., 28: 59-65, l992

Burstein, M. and D. Wool. Gall aphids do not select optimal sites (Smynthurodes betae, Pemphigidae). Ecol. Entomol. 18: 155-164, 1993.

Kamin-Belsky, N. and D. Wool. Ecological aspects of digestive enzyme co-variation in almond moth (Ephestia cautella (Walker) (Lepidoptera, Phyticidae). J. Stored Prod. Res. 29: 323-332, 1993.

Wool, D., D. Gerling, A.C. Bellotti and F.J. Morales. Esterase electrophoretic variation in Bemisia tabaci (Genn.) (Hom. Aleyrodidae) among host plants and localities in Israel. J. Appl. Ent. 115: 185-196, 1993.

Shchukin, A. and D. Wool. Pyrethroid resistance and esterase activity in selected laboratory populations of sweetpotato whiteflies (Bemisia tabaci). Eur. J. Entomol. 91: 285-295, 1994.

Bloch, G. and D. Wool. Methidathion resistance in the sweetpotato whitefly Bemisia tabaci (Aleyrodidae: Homoptera) in Israel: selection, heritability, and correlated changes of esterase activity. J. Econ. Entomol. 87: 1147-1156, 1994.

Inbar (Burstein), M. and D. Wool. Phloem-feeding specialists sharing a host tree: resource partitioning minimizes
interference competition among galling aphid species. Oikos 73: 109-119, 1994.


Burstein, M., D. Wool and A. Eshel. Sink strength and clone size of sympatric gall-forming aphids. Eur. J. Entomol. 91: 57-61, 1994.

Wool, D., O. Manheim, M. Burstein and T. Levi. Dynamics of re-migration of sexuparae to their primary hosts in the gall-forming Fordinae (Aphidoidea: Pemphigidae). Eur. J. Entomol. 91: 103-108, 1994.

Wool, D., L. Calvert, L.M. Constantino, A.C. Bellotti and D. Gerling. Differentiation of Bemisia tabaci (Genn.) populations in Colombia. J. Appl. Ent. 117: 122-134, 1994.

Wool, D. Aphid-induced galls on Pistacia in the natural Mediterranean forest of Israel: which, where, and how many? Isr. J. Zool. 41: 591-600, 1995.

Bloch, G. and D. Wool. Esterase activity in populations of the whitefly, Bemisia tabaci: heritability and associated organophosphorous resistance. Bull. ent. Res. 85: 11-19, 1995.

Wool, D. and N. Bar-El. Population ecology of the galling aphid Forda formicaria von Heyden in Israel: abundance, demography and gall structure. Isr. J. Zool. 41: 175-192, 1995.

Inbar, M., A. Eshel and D. Wool. Interspecific competition among phloem-feeding insects mediated by induced hostplant sinks. Ecology 76: 1506-1515, 1995.

Wool, D., D. Hales and P. Sunnucks. Host plant relationships of Aphis gossypii Glover (Homoptera: Aphididae) in Australia. J. Austral. Ent. Soc. 34: 265-271, 1995.

Wool, D. Aphid galls: what makes them attractive for study? Cecidology 11: 2-8, 1996.

Wool, D. and D.F. Hales. Previous infestation affects recolonization of cotton by Aphis gossypii: induced resistance or plant damage?
Phytoparasitica 24: 39-48, 1996.

Wool, D. and D.F. Hales. Phenotypic plasticity in Australian Aphis gossypii: hostplant effects on morpholoigcal variation. Ann. Ent. Soc. Amer. 90: 316-328, 1997.

Wool, D., O. Manheim and M. Inbar. The return flight of sexuparae of galling aphids to their primary host trees: implications for differential herbivory and gall abundance (Aphidoidea: Pemphigidae, Fordinae). Ann. Ent. Soc. Amer. 90: 341-350, 1997.

Yom Tov, Y. and D. Wool. Do the contents of Barn Owl pellets accurately represent the proportion of prey species in the field? Condor 99: 972-976, 1997.


Wool, D. and M. Inbar. Colonization of ecological islands: galling aphid populations on recovering Pistacia trees after destruction by fire. Eur. J. Entomol. 95: 41-53, 1998

Wool ,D., and O. Ben Zvi Population Ecology and clone dynamics of the galling aphid Geoica wertheimae (Sternorrhynch: Pemphigidae: Fordinae) Eur.J.Entomol. 95: 509-518, 1998

Wool, D. and R. Bogen Ecology of the gall-forming aphid, Slavum wertheimae, On Pistacia atlantica : population dynamics and differential herbivory. Isr.J.Zool. 45: 247-260, 1999

Wool, D., R. Aloni, O. Ben Zvi and M. Wollberg A galling aphid furnishes its home with a built-in pipeline to the host food supply.
Entomol. Exp. Appl. 91:183-186, 1999

Wool, D. and T. Agami Response to selection and genetic regulation of esterase activity variation in Tribolium confusum
Selection 1: 95-105, 2000

Wool, D. Charles Lyell - " The father of Geology" - as a fore runner of modern Ecology Oikos 94: 385-391, 2001

Wool, D. and Sulami, Z. Induction of alate sexuparae in root cage colonies, and female-biased sex ratios in the galling aphid, Aploneura lentisci Entomol. Exp. Appl. 101: 299-303, 2001

Wool, D. Herbivore abundance is independent of weather? A 20-year study of a galling aphid, Baizongia pistaciae (Homoptera: Aphidoidea.) Popul. Ecol. 44: 281-291, 2002


Dayan, T., Wool, D., and Simbersloff, D. Variation and covariation of skulls and teeth: modern carnivores and the interpretation of fossil mammals Paleobiology 28: 508-526, 2002


Wool, D., and Front, L. Esterase variation in Tribolium confusum (Coleoptera: Tenebrionidae) : genetic analysis of interstrain crosses in relation to malathion resistance J. Stored Prod. Res. 39: 237-249, 2003

Martinez, J.J.(Y.) and Wool, D. Differential response of trees and shrubs to browsing and pruning : the effect on Pistacia growth and gall-inducing aphids Plant Ecology 169: 285- 294, 2003


Wool, D. Galling aphids : Specialization, biological complexity, and variation. Annu. Rev. Entomol. 49: 175- 194, 2004

Manheim, O., and Wool, D. Ifferential response of genotypes to alternative environments: a comparative morphological study of gall-inducing aphids ( Homoptera: Pemphigidae: Fordinae) Israel J.Zool. 49: 287-305, 2003

Inbar, M., Wink M. and Wool, D. The evolution of host-plant manipulation by insects: molecular and ecological evidence from gall-forming aphids on Pistacia Molecular Phylogenetics and Evolution, 2004


Martinez, J-J.Y. , Mokady O., Wool D. Patch size and patch quality of gall- inducing aphids in a mosaic landscape in Israel. Landscape Ecology, 20:1013-1024, 2005.


Hazkani-Covo,E. Wool, D., Graur, D.  In search of the vertebrate phylotypic stage: a molecular examination of the development hourglass model and Von Baer's third law. J. Exp. Zool. (Mol.Dev.Evol.) 304 B: 150-158 , 2005.


Wool, D. Differential colonization of host trees by galling aphids: selection of hosts or selection by hosts? Basic and Applied Ecology, 6: 445-451, 2005.


Wool, D., Hendrix, D.L., Shukry, O. Seasonal variation in honeydew sugar content of galling aphids ( Aphidoidea: Pemphigidae: Fordinae) feeding on Pistacia : Host ecology and aphid physiology. Basic and Applied Ecology, 7: 141-151


Martinez, J-J. Y. , Wool, D. Sampling bias in roadsides: the case of galling aphids on Pistacia trees Biodiversity and Conservation, 2006 DOI 10.1007/s 10531-004-6685-2



Chapters in books

Wool, D. and J. Koach. Morphological variation of the gall-forming Aphid, Geoica utricularia (Homoptera) in relation to environmental variation. pp. 239-272 In: "Population Genetics and Ecology", S. Karlin and E. Nevo, (eds.). Academic Press, 1976.


Wool, D. Gall-forming Aphids. pp.l1-58 In: "Biology of gall insects", T.N. Ananthakrishnan (ed.). Oxford & IBH, 1984.


Wool, D. Electrophoretic variation in post-harvest agricultural pests, and its implications. pp. 341-362 In: "Electrophoretic studies on agricultural pests", H.D. Loxdale and J. Den Hollander (eds.). Clarendon Press, Oxford, 1989.


Wool, D. and O. Manheim. Multivariate study of temporal variation in morphology of a gall-forming aphid. pp. 179-192 In: "Ordination in the study of morphology, evolution and systematics of insects", J. Sorensen and R. Foottit (eds.). Elsevier, 1992.


Wool, D. The shapes of insect galls: insect control, plant constraints and phylogeny. pp. 203-213 In: "Ecology and evolution of plant-feeding insects in natural and man-made environments", A. Raman (ed.). Intern. Sci. Publ., New Delhi, India, 1997.


Wool, D. Gall-forming aphids on Pistacia: a first look at the subterranean part of their life cycle. pp.127-131 in “The biology of gall-inducing arthropods”, G.Csoka, W.J.Mattson, G.N.Stone and P.W.Price (eds.), USDA Forest Service , St. Paul, Minnesota, 1998.

Wool, D. Gall-inducing aphids : biology, ecology and evolution. In : "Biology, Ecology and Evolution of Gall-inducing Arthropods"; A. Raman, C.W.Schaefer, and T.M. Withers,(eds.). Science Publishers, New Hempshire, USA.


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