Adam Tofilski

Department of Zoology and Animal Welfare
Agricultural University
29 Listopada 54
31-425 Krakow

Tel: +48 12 6625069

polska wersja

More about me:
Google Scholar

My software:
IdentiFly - for insect identification based on geometric morphometrics
DKey - for editing and browsing dichotomous taxonomic keys

My web pages:
Honey bee - some information about honey bee biology
DrawWing - some information about insect wings

My publications:

Puškadija Z., Kovačić M., Raguž N., Lukić B., Prešern J., Tofilski A. 2021. Morphological diversity of Carniolan honey bee (Apis mellifera carnica) in Croatia. Journal of Apicultural Research, (in press)

Janczyk A., Meixner M.D., Tofilski A. 2021. Morphometric identification of the endemic Maltese honey bee (Apis mellifera ruttneri). Journal of Apicultural Research. 60: 157-164.

Hailu, T. G., D’Alvise, P., Tofilski A., Fuchs, S., Greiling, J., Rosenkranz, P., Hasselmann, M. (2020). Insights into Ethiopian honey bee diversity based on wing geomorphometric and mitochondrial DNA analyses. Apidologie 51: 1182-1198.

Goczał J., Oleksa A., Rossa R., Chybicki I., Meyza K., Plewa R., Landvik M., Gobbi M., Hoch G., Tamutis V., Balalaikins M., Telnov D., Dascalu M., Tofilski A. 2020. Climatic oscillations drive plants–insects co–evolution: Divergence patterns across the distribution range of Monochamus sartor reflect Quaternary history of its host tree. Scientific Reports 10: 16524.

Czekońska K., Tofilski A. 2020. Body mass of honey bee drones developing in constant and in changing temperatures. Apidologie, 51: 510–518.

Łopuch, S., Tofilski A. 2020. Impact of the quality of food sources on the wing beating of honey bee dancers. Apidologie 51: 631–641.

Czekońska K., Szentgyörgyi H., Tofilski A. 2019. Body mass but not wing size or symmetry correlates with life span of honey bee drones. Bulletin of Entomological Research 109: 383–389.

Łopuch, S., Tofilski A. 2019. Use of high-speed video recording to detect wing beating produced by honey bees. Insectes Sociaux, 66: 235–244.

Szpila K., Żmuda A., Akbarzadeh K., Tofilski A. 2019. Wing measurement can be used to identify European blow flies (Diptera: Calliphoridae) of forensic importance. Forensic Science International, 296: 1–8.

Goczał J., Rossa R., Tofilski A., 2019. Intersexual and intrasexual patterns of horn size and shape variation in the European rhinoceros beetle: quantifying the shape of weapons. Biological Journal of the Linnean Society, 127: 34-43.

Węgrzynowicz P., Gerula D., Tofilski A., Panasiuk B., Bieńkowska M., 2019. Maternal inheritance in hybrids of three honey bee subspecies. Journal of Apicultural Science, 63: 131-138.

Szentgyörgyi H., Czekońska K., Tofilski A. 2018 Honey bees are larger and live longer after developing at low temperature. Journal of Thermal Biology 78: 219-226.

Tofilski A., Chuda-Mickiewicz B., Czekońska K. 2018. Circular movement of honey bee spermatozoa inside spermatheca. Invertebrate Reproduction & Development 62: 63-66.

Goczał J., Rossa R., Nawrocka A., Sweeney J.D., Tofilski A. 2018. Developmental costs of biological invasion: The exotic wood borer Tetropium fuscum (Coleoptera: Cerambycidae) is more asymmetric and smaller in invaded area. Environmental Entomology 47: 982–989.

Mielczarek Ł.E., Tofilski A. 2018. Semiautomated identification of a large number of hoverfly (Diptera: Syrphidae) species based on wing measurements. Oriental Insects 52: 245-258.

Nawrocka A., Kandemir I., Fuchs S., Tofilski A. 2018. Computer software for identification of honey bee subspecies and evolutionary lineages. Apidologie 49: 172-184.

Goczał J., Rossa R., Tofilski A. 2018. Elytra reduction may affect the evolution of beetle hind wings. Zoomorphology 137: 131-138.

Tofilski A. 2018. DKey software for editing and browsing dichotomous keys. ZooKeys 735: 131-140.

Łopuch S., Tofilski A. 2017. Importance of wing movements for information transfer during honey bee waggle dance. Ethology 123: 974–980.

Szentgyorgyi H., Moroń D., Nawrocka A., Tofilski A., Woyciechowski M. 2017. Forewing structure of the solitary bee Osmia bicornis developing on heavy metal pollution gradient. Ecotoxicology 26: 1031-1040.

Szentgyörgyi H., Czekońska K., Tofilski A. 2017. The effects of starvation of honey bee larvae on reproductive quality and wing asymmetry of honey bee drones. Journal of Apicultural Science 61: 233-243.

Goczał J., Rossa R., Sweeney J., Tofilski A. 2017. Citizen monitoring of invasive species: wing morphometry as a tool for detection of alien Tetropium species. Journal of Applied Entomology 141: 496-506.

Grzywacz A., Ogiela J., Tofilski A. 2017. Identification of Muscidae (Diptera) of medico-legal importance by means of wing measurements. Parasitology Research 116: 1495-1504.

Łopuch S., Tofilski A. 2017. Direct visual observation of wing movements during the honey bee waggle dance. Journal of Insect Behavior 30: 199-210.

Rossa R., Goczał J., Tofilski A. 2017. Hind wing morphology facilitates discrimination between two sibling species: Leiopus nebulosus and L. linnei (Coleoptera: Cerambycidae). Zootaxa 4227: 266–278.

Szentgyörgyi H., Czekońska K., Tofilski A. 2016. Influence of pollen deprivation on the fore wing asymmetry of honeybee workers and drones. Apidologie 47:653-662.

Łopuch S., Tofilski A. 2016. The relationship between asymmetry, size and unusual venation in honey bees (Apis mellifera). Bulletin of Entomological Research 106:304–313.

Przybyłowicz Ł., Pniak M., Tofilski A. 2016. Semiautomated Identification of European Corn Borer (Lepidoptera: Crambidae). Journal of Economic Entomology 109:195-199.

Rossa R., Goczał J., Tofilski A. 2016. Within- and between-species variation of wing venation in genus Monochamus (Coleoptera: Cerambycidae). Journal of Insect Science 16:5.

Mielczarek L.E., Oleksa A., Meyza K., Tofilski A. 2016. Seasonal polyphenism in Eristalis pertinax (Diptera: Syrphidae). European Journal of Entomology 113:489–496.

Oleksa A., Tofilski A. 2015. Wing geometric morphometrics and microsatellite analysis provide similar discrimination of honey bee subspecies. Apidologie 46:49-60.

Tofilski A. 2014. A scientific note on amoeboid movement of honey bee semen. Apidologie 45:637-640.

Soszyński B., Mielczarek L.E., Tofilski A. 2013. Dasysyrphus neovenustus sp. n. (Diptera: Syrphidae) a new species in the venustus species group. Polish Journal of Entomology 82:353-363.

Wenseleers T., Bacon J.P., Alves D.A., Couvillon M.J., Kärcher M., Nascimento F.S., Nogueira-Neto P., Ribeiro M., Robinson E.J.H., Tofilski A., Ratnieks F.L.W. 2013. Bourgeois behavior and freeloading in the colonial orb web spider Parawixia bistriata (Araneae, Araneidae). American Naturalist 182:120-129.

Oleksa A., Wilde J., Tofilski A., Chybicki I.J. 2013. Partial reproductive isolation between European subspecies of honey bees. Apidologie 44:611-619.

Tofilski A., Oleksa A. 2013. There are still bee trees in Europe. Bee World 90:18-19.

Oleksa A., Gawroński R., Tofilski A. (2013) Rural avenues as a refuge for feral honey bee population. Journal of Insect Conservation 17:465-472. Available online

Tofilski A., Chuda-Mickiewicz B., Czekonska K., Chorbiński P. (2012) Flow cytometry evidence about sperm competition in honey bee (Apis mellifera). Apidologie 43:63-70.

Oleksa A., Chybicki I., Tofilski A., Burczyk J., 2011. Nuclear and mitochondrial patterns of introgression into native dark bees (Apis mellifera mellifera) in Poland. Journal of Apicultural Research 50:116-129.

Bonckaert W., Tofilski A., Nascimento F., Billen J., Ratnieks F., Wenseleers T., 2011. Co-occurrence of three types of egg policing in the Norwegian wasp Dolichovespula norwegica. Behavioral Ecology and Sociobiology 65:633-640.

Tofilski A., 2011. Homology of submarginal cross veins in forewings of bees (Hymenoptera: Apiformes). Journal of Apicultural Science 55:131-140.

Tofilski A., 2009. Shorter-lived workers start foraging earlier. Insectes Sociaux 56:359-366.

Gerula D., Tofilski A., Węgrzynowicz P., Skowronek W., 2009. Computer-assisted discrimination of honeybee subspecies used for breeding in Poland. Journal of Apicultural Science 53:105-114.

Tofilski, A., M. J. Couvillon, S. E. F. Evison, H. Helanterä, E. J. H. Robinson, and F. L. W. Ratnieks. 2008. Pre-emptive defensive self-sacrifice by ant workers. American Naturalist 172:E239-E243.

Tofilski A., 2008. Using geometric morphometrics and standard morphometry to discriminate three honeybee subspecies. Apidologie 39:558-563.

Tofilski, A., Ratnieks F.L.W. 2008. Simple rules based on pile slope are used in the self organization of sand pile formation by Pheidole oxyops ants. Insectes Sociaux 55:37-42.

Robinson E. J. H., Tofilski A., Ratnieks F. L. W., 2008. The use of native and non-native tree species for foraging and nesting habitat by the wood-ant Formica lugubris (Hymenoptera: Formicidae). Myrmecological News 11:1-7.

Tofilski, A. 2007. Automatic measurements of honeybee wings. In MacLeod, N. (ed.), Automated Object Identification in Systematics: Theory, Approaches, and Applications, Systematics Association Special Volume, CRC Press, Boca Raton, Florida, pp 289-298.

Tofilski A. 2006. Influence of caste polyethism on longevity of workers in social insect colonies. J. Theor. Biol. 238:527-531.
PDF, Appendix, Mathematica 4.0 notebook

D'Ettorre P., Tofilski A., Heinze J., Ratnieks F. L. W., 2006. Non-transferable signals on ant queen eggs. Naturwissenschaften 93:136-140.

Helanterä H., Tofilski A., Wenseleers T., Ratnieks F. L. W., 2006. Worker policing in the common wasp Vespula vulgaris is not aimed at improving colony hygiene. Insectes Sociaux 53:399-402.

Tofilski A., Ratnieks F.L.W. 2005. Sand pile formation in Dorymyrmex ants. Journal of Insect Behavior 18:505-512.

Wenseleers T, Tofilski A, Ratnieks FLW, 2005. Queen and worker policing in the tree wasp Dolichovespula sylvestris. Behavioral Ecology and Sociobiology 58:80-86.

Wenseleers T, Badcock NS, Erven K, Tofilski A, Nascimento FS, Hart AG, Burke TA, Archer ME, Ratnieks FLW, 2005. A test of worker policing theory in an advanced eusocial wasp, Vespula rufa. Evolution 59:1306-1314.

Nascimento F. S., Hrncir M., Tofilski A., Zucchi R., 2005. Scraping sounds produced by a social wasp (Asteloeca ujhelyii, Hymenoptera: Vespidae). Ethology 111:1116-1125.

Jensen A., Palmer K., Chaline N., Raine N., Tofilski A., Martin S., Pedersen B., Boomsma J., Ratnieks F., 2005. Quantifying honey bee mating range and isolation in semi-isolated valleys by DNA microsatellite paternity analysis. Conservation Genetics 6:527-537.

Tofilski A. 2004. DrawWing, a program for numerical description of insect wings. Journal of Insect Science, 4:17, 5pp.
Available online:

Tofilski A., Czekonska K. 2004. Emergency queen rearing in honeybee colonies with brood of known age. Apidologie 35:275-282.

Tofilski A. 2002. Influence of age polyethism on longevity of workers in social insects. Behav. Ecol. Sociobiol. 51: 234-237.
PDF , Appendix, Mathematica 4.0 notebook

Tofilski A. 2000. Senescence and learning in honey bee (Apis mellifera) workers. Acta. Neurobiol. Exp. 60:35-39.

Tofilski A., Kopel J. 1996. The influence of Nosema apis on maturation and flight activity of honey bee drones. Pszczelnicze Zeszyty Naukowe 40:55-60.

Tofilski A. 1995. When should honey bee (Apis mellifera) colonies swarm? I Krajowa Konferencja Zastosowañ Matematyki w Biologii i Medycynie, Zakopane.

Last Updated 2021-02-08