Department of Animal Ecology and Tropical Biology

Prof. Dr. Thomas Schmitt

Telephone: +49 (0)931 31-84188
Fax:             +49 (0)931 31-84352
e-Mail:  thomas.schmitt@uni-wuerzburg.de
ORCID: 0000-0002-6719-8635

Room C016

University of Würzburg
Department of Animal Ecology and Tropical Biology
Biocenter -  Am Hubland

97074 Würzburg, Germany

  • Evolution of chemical communication in insects

  • Evolution of cuticular hydrocarbon profiles in insects

  • Functional aspects of chemical interactions in animals and plants

  • since 2013  Professor of Animal Ecology at the University of Würzburg
  • 2012 – 2013  Research Associate at the Technical University of Darmstadt
  • 2010  Habilitation in Evolutionary Biology and Animal Ecology at the University of Freiburg
  • 2004 – 2012  Research Associate at the University of Freiburg
  • 2004  Doctoral thesis: „Communication in the Hymenoptera: Chemistry, Ecology, and Evolution“ at the University of Würzburg
  • 2000  Diploma thesis: „Chemical ecology and niche specialization in West African dung beetles” at the University of Würzburg
  • 1994 – 2000  Study of Biology at the University of Würzburg and SUNY Albany, NY, USA
  • 1992 – 1994  Study of Chemistry at the University of Würzburg

[in press]




Ala-Honkola O, Kauranen H, Tyukmaeva V, Boetzl FA, Hoikkala A, Schmitt T (2020) Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana. Insect Science 27: 304-316. https://doi.org/10.1111/1744-7917.12639.

Polidori C, Ballesteros Y, Wurdack M, Asís JD, Tormos J, Baños-Picón L, Schmitt T (2020) Low host specialization in the cuckoo wasp, Parnopes grandior, weakens chemical mimicry but does not lead to local adaption. Insects 11: 136. https://doi.org/10.3390/insects11020136. OPEN ACCESS.

Polidori C, Geyer M, Schmitt T (2020) Do Sphecodes cuckoo bees use chemical insignificance to invade the nests of their social Lasioglossum bee hosts? Apidologie 51, 147-162.  https://doi.org/10.1007/s13592-019-00692-x.


Drescher N, Klein A-M, Schmitt T, Leonhardt SD (2019) A clue on bee glue: New insight into the sources and factors driving resin intake in honeybees (Apis mellifera). PLoS ONE 14 (2): e0210594. https://doi.org/10.1371/journal.pone.0210594. OPEN ACCESS.

Hartke J, Schell T, Jongepier E, Schmidt H, Sprenger PP, Paule  J, Bornberg-Bauer E, Schmitt T, Menzel F, Pfenninger M, Feldmeyer B (2019) Hybrid genome assembly of a neotropical mutualistic ant. Genome Biology and Evolution 11: 2306-2311. https://doi.org/10.1093/gbe/evz159.

Hartke J, Sprenger PP, Sahm J, Winterberg H, Orivel J, Baur H, Beuerle T, Schmitt T, Feldmeyer B, Menzel F (2019) Cuticular hydrocarbons as potential mediators of cryptic species divergence in a mutualistic ant association. Ecology and Evolution 9: 9160-9176. https://doi.org/10.1002/ece3.5464. OPEN ACCESS.

Kämper W, Kaluza BF, Wallace H, Schmitt T, Leonhardt SD (2019) Habitats shape the cuticular chemical profiles of stingless bees. Chemoecology 29:125-133. https://doi.org/10.1007/s00049-019-00282-4

Pauli T, Castillo-Cajas RF, Rosa P, Kukowka S, Berg A, van den Berghe E, Fornoff F, Hopfenmüller S, Niehuis N, Peters RS, Staab M, Strumia F, Tischendorf S, Schmitt T,  Niehuis O (2019) Phylogenetic analysis of cuckoo wasps (Hymenoptera: Chrysididae) reveals a partially artificial classification at the genus level and a species-rich clade of bee parasitoids. Systematic Entomology44: 322-335. doi.org/10.1111/syen.12323.

Poikela N, Kinnunen J, Wurdack M, Kauranen H, Schmitt T, Kankare M. Snook RR, Hoikkala A (2019) Strength of sexual and postmating prezygotic barriers varies between sympatric populations with different histories and species abundances. Evolution  73: 1182–1199. doi.org/10.1111/evo.13732.

Sprenger PP, Hartke J, Feldmeyer B, Orivel J, Schmitt T, Menzel F (2019) Influence of mutualistic lifestyle, mutualistic partner, and climate on cuticular hydrocarbon profiles in parabiotic ants. Journal of Chemical Ecology 45: 741-754. doi.org/10.1007/s10886-019-01099-9.

Streinzer M, Chakravorty J, Neumayer J, Megu K, Narah J, Schmitt T, Bhardi H, Spaethe J, Brockmann A (2019) Species composition and elevational distribution of bumble bees (Hymenoptera, Apidae, Bombus Latreille) in the East Himalaya, Arunachal Pradesh, India. ZooKeys 851: 71-89. doi.org/10.3897/zookeys.851.32956.


Büllesbach J, Cash E, Schmitt T (2018) Communication, Insects. IN: Skinner MK (ed), Encylopedia of Reproduction. Vol 6, pp 78-83. https://doi.org/10.1016/B978-0-12-809633-8.20539-6.

Büllesbach J, Vetter SG, Schmitt T (2018) Differences in the reliance on cuticular hydrocarbons as sexual signaling and species discrimination cues in parasitoid wasps. Frontiers in Zoology 15: 22. https://doi.org/10.1186/s12983-018-0263-z . OPEN ACCESS.

Frank K, Brückner A. Blüthgen N, Schmitt T (2018) In search of cues: dung beetle attraction and the significance of volatile composition of dung. Chemoecology 28: 145-152. https://doi.org/10.1007/s00049-018-0266-4.

Frank K, Krell FT, Slade EM, Raine EH, Chiew LY, Schmitt T, Vairappan CS, Walter P, Blüthgen N (2018) Global dung webs: high trophic generalism of dung beetles along the latitudinal diversity gradient. Ecology Letters 21: 1229-1236. https://doi.org/10.1111/ele.13095.

Pull CD, Ugelvig LV, Wiesenhofer F, Grasse AV, Tragust S, Schmitt T, Brown MJF, Cremer S (2018) Destructive disinfection of infected brood prevents systemic disease spread in ant colonies. Elife 7: e32073. https://doi.org/10.7554/eLife.32073.001. OPEN ACCESS


Bank S., Sann M., Mayer C., Meusemann K., Donath A., Podsiadlowski L., Kozlov A., Petersen M., Krogmann L., Meier R., Rosa P., Schmitt T., Wurdack M., Liu S., Zhou X., Misof B., Peters R. S., Niehuis O. (2017) Transcriptome and target DNA enrichment sequence data provide new insights into the phylogeny of vespid wasps (Hymenoptera: Aculeata: Vespidae). Molecular Phylogenetics and Evolution 116: 213–226. https://doi.org/10.1016/j.ympev.2017.08.020.

Frank K, Hülsmann M, Assmann T, Schmitt T,  Blüthgen N (2017) Land use affects dung beetle communities and their ecosystem service in forests and grasslands. Agriculture, Ecosystems and Environment 243: 114-122. https://doi.org/10.1016/j.agee.2017.04.010.

Frank ET, Schmitt T,  Hovestadt T, Mitesser O, Stiegler J, Linsenmair KE (2017) Saving the injured: Rescue behavior in the termite-hunting ant Megaponera analis. Science Advances 3, e1602187. https://doi.org/10.1126/sciadv.1602187.

Haberer W, Schmitt T, Schreier P, Eggert A-K, Müller JK (2017) Volatiles emitted by calling males of burying beetles and Ptomascopus morio (Coleoptera: Silphidae: Nicrophorinae) are biogenetically related. Journal of Chemical Ecology 43: 971-977. https://doi.org/10.1007/s10886-017-0892-2.

Junker RR, Kuppler J, Amo L, Blande JD, Borges RM, van Dam NM, Dicke M, Dötterl S, Ehlers B, Etl F, Gershenzon J, Glinwood R, Gols R, Groot AT, Heil M, Hoffmeister M, Holopainen JK, Jarau S, John L, Kessler A, Knudsen JT, Kost C, Larue-Kontic AAC, Leonhardt SD, Lucas-Barbosa D, Majetic CJ, Menzel F, Parachnowitsch AL, Pasquet RS, Poelman EH, Raguso RA, Ruther J, Schiestl FP, Schmitt T, Tholl D, Unsicker SB, Verhulst N, Visser ME, Weldegergis BT, Köllner TG (2017) Co-variation and phenotypic integration in chemical communication displays: biosynthetic constraints and eco-evolutionary implications. New Phytologist 220: 739-749. https://doi.org/10.1111/nph.14505.

Menzel F, Schmitt T, Blaimer B (2017) The evolution of a complex trait: cuticular hydrocarbons in ants evolve independently from phylogenetic constraints. Journal of Evolutionary Biology 30, 1372-1385. https://doi.org/10.1111/jeb.13115.

Menzel F, Blaimer B, Schmitt T (2017) How do cuticular hydrocarbons evolve? Physiological constraints, climatic and biotic selection pressures act on a complex functional trait in insects. Proceedings of the Royal Society B, London 284, 20161727. https://doi.org/10.1098/rspb.2016.1727.

Peters RS, Krogmann L, Mayer C, Donath A, Gunkel S, Meusemann K, Kozlov A, Podsiadlowski L, Petersen M, Lanfear R, Diez PA, Heraty J, Kjer KM, Klopfstein S, Meier R, Polidori C, Schmitt T, Liu S, Zhou X, Wappler T, Rust J, Misof B, Niehuis O (2017) Evolutionary History of the Hymenoptera. Current Biology 27, 1013-1018. https://doi.org/10.1016/j.cub.2017.01.027.

Polidori C, Giordani I, Wurdack M, Tormos J, Asís JD, Schmitt T (2017) Post-mating shift towards longer-chain cuticular hydrocarbons drastically reduces female attractiveness to males in a digger wasp. Journal of Insect Physiology 100, 119-127. https://doi.org/10.1016/j.jinsphys.2017.05.001.

Wurmitzer C, Blüthgen N, Krell FT, Maldonado B, Ocampo F, Müller JK, Schmitt T (2017) Attraction of dung beetles to herbivore dung and synthetic compounds in comparative field study. Chemoecology 27, 75-84. https://doi.org/10.1007/s00049-017-0232-6.

Wurdack M, Polidori C, Keller A, Feldhaar H, Schmitt T  (2017) Release from prey preservation behavior via prey switch allowed diversification of cuticular hydrocarbon profiles in digger wasps. Evolution 71: 2562-2571. https://doi.org/10.1111/evo.13322.



Flaig I. C., Aguilar I., Schmitt T. & Jarau S. (2016) An unusual recruitment strategy in a mass‐recruiting stingless bee, Partamona orizabaensisJournal of Comparative Physiology A 202: 679-690.

Diao W., Mousset M., Horsburgh G. J., Vermeulen C. J., Johannes F., van de Zande L., Ritchie M. G., Schmitt T. & Beukeboom L. W. (2016) Quantitative trait locus analysis of mating behavior and male sex pheromones in Nasonia wasps. G3 - Genes, Genomics, Genetics 6: 1549-1562.

Leonhardt S. D., Menzel F., Nehring V. & Schmitt T. (2016) Ecology and evolution of communication in social insects (invited review). Cell 164: 1277-1287.
Jansen J., Pokorny T. & Schmitt T. (2016): Disentangling the effect of insemination and ovary development on the cuticular hydrocarbon profile in the bumblebee Bombus terrestris (Hymenoptera: Apidae). Apidologie 47: 101-113.


Wurdack M., Herbertz S., Dowling D., Kroiss J., Strohm E., Baur H., Niehuis O. & Schmitt T. (2015) Striking cuticular hydrocarbon dimorphism in the mason wasp Odynerus spinipes and its possible evolutionary cause (Hymenoptera: Chrysididae, Vespidae). Proceedings of the Royal Sociecty B, London 282: 20151777.

Strube-Bloss M. F., Brown A., Spaethe J., Schmitt T. & Rössler W. (2015) Extracting the behaviorally relevant stimulus: Unique neural representation of farnesol, a component of the recruitment pheromone of Bombus terrestris. PLoS ONE 10: e0137413. OPEN ACCESS.

Buellesbach J. & Schmitt T. (2015): The role of cuticular hydrocarbon-based chemical communication in prezygotic reproductive isolation. In Animal Communication and Cognition: Principles, Evolution and Development, ed. Wagner T. Nova Publishers, Hauppauge NY, pp. 107-119.


Leonhardt S. D., Baumann A.-M., Wallace H. M., Brooks P. & Schmitt T. (2014) The chemistry of an unusual seed dispersal mutualism: bees use a complex set of olfactory cues to find their partner. Animal Behaviour 98: 41-51.

Büllesbach J., Greim C., Raychoudhury R. & Schmitt T. (2014) Asymmetric assortative mating behaviour reflects incomplete prezygotic isolation in the Nasonia species complex. Ethology 120: 834-843.

Jennings J. H., Etges W. J., Schmitt T. & Hoikkala A. (2014) Cuticular hydrocarbons of Drosophila montana: Geographic variation, sexual dimorphism and potential roles as pheromones. Journal of Insect Physiology 61: 16–24.

Menzel F., Orivel J., Kaltenpoth M. & Schmitt T. (2014) What makes you a potential partner? Insights from convergently evolved ant–ant symbioses. Chemoecology 24: 105-119.


Giesbers M. C. W. G., Gerritsma S., Buellesbach J., Diao W., Pannebaker B. A., van de Zande L., Schmitt T. & Beukeboom L. W. (2013): Prezygotic isolation in the parasitoid wasp genus Nasonia. In: Speciation: Natural Processes, Genetics and Biodiversity, ed. Michalak P.  Nova Publishers, Hauppauge NY, pp. 165-195.

Rödel M.-O., Brede C., Hirschfeld M., Schmitt T., Favreau P., Stöcklin R., Wunder C. & Mebs D. (2013) Chemical camouflage – a frog’s strategy to co-exist with aggressive ants. PLOS One 8(12):e81950.

Buellesbach J. M., Gadau J., Beukeboom L. W., Echinger F., Raychoudhury R., Werren J. H. & Schmitt T. (2013) Cuticular hydrocarbon divergence in the jewel wasp Nasonia: Evolutionary shifts in chemical communication channel? Journal of Evolutionary Biology 26: 2467-2478.

Menzel F., Blüthgen N., Tolasch T., Conrad J., Beifuß U., Beuerle T. & Schmitt T. (2013) Crematoenones – a novel substance class exhibited by ants functions as appeasement signal. Frontiers in Zoology 10:32.

Leonhardt S. D., Rasmussen C. & Schmitt T. (2013) Genes vs. environment: Geography and phylogenetic relationships shape the chemical profiles of stingless bees on a global scale. Proceedings of the Royal Society B, London 280:1762.

Niehaus O., Büllesbach J., Gibson, J. D., Pothmann D., Hanner C., Mutti N. S., Judson A. K., Gadau J., Ruther J. & Schmitt T. (2013) Behavioural and genetic analyses on Nasonia shed light on the evolution of sex pheromones. Nature 494: 345-348.


Schlechter J., Schmitt T. & Peschke K. (2012) Learning individual signatures: rove beetle males discriminate unreceptive females by cuticular hydrocarbon patterns. Animal Behaviour 84: 369-376.

Menzel F. & Schmitt T. (2012) First evidence for an interspecific selection pressure on chemical recognition cues. Evolution 66: 896-904.

Butlin R., Debelle A., Kerth C, Snook R. R., Beukeboom L. W., Castillo Cajas R. F., Diao W., Maan M. M., Paolucci S., Weissing F. J., van de Zande L., Hoikkala A., Geuverink E., Jennings J., Kankare, M., Knott K. E., Tyukmaeva V. I., Zoumadakis C., Ritchie M. G., Barker D., Immonen E., Noor M., Macias C., Schmitt T. & Schilthuizen M. (2012) What do we need to know about speciation? Trends in Ecology and Evolution 27: 27-39.


Leonhardt S.D., Form S., Blüthgen N., Schmitt T. & Feldhaar H. (2011) Genetic relatedness and chemical profiles in an unusually peaceful eusocial bee. Journal of Chemical Ecology 37: 1117-1126.

Schlechter J., Schmitt T. & Peschke K. (2011) A contact antiaphrodisiac pheromone supplied by the spermatophore in the rove beetle Aleochara curtula: mode of transfer and evolutionary significance. Naturwissenschaften 98: 855-862.

Haberer W., Schmitt T., Schreier P. & Müller J.K. (2011) Intended and unintended receivers of the male pheromones of the burying beetles Nicrophorus humator and Nicrophorus vespilloidesEntomologia Experimentalis et Applicata 140: 122-126.

Leonhardt S., Schmitt T. & Blüthgen N. (2011) Tree resin composition, collection behaviour and selective filters shape chemical profiles of tropical bees (Apidae: Meliponini). PLoS One 6(8), e23445.

Leonhardt, S.D., Wallace, H.M. & Schmitt T. (2011) The cuticular profiles of Australian stingless bees are shaped by resin of the euclypt tree Corymbia torelliana. Austral Ecology 36: 537-543.

Niehuis O., Büllesbach J., Judson A.K., Schmitt T. & Gadau J. (2011) Genetics of cuticular hydrocarbons in F2 hybrid males of Nasonia giraulti and Nasonia vitripennisHeredity 107: 61-70.

Steiger S., Schmitt T. & Schaefer H. M. (2011) The dynamic evolution of chemical information transfer. Proceedings of the Royal Society B, London 278: 970-979.

Leonhardt S., Blüthgen N. & Schmitt T. (2011) Chemical profiles of body surfaces and nests from six Bornean stingless bee species. Journal of Chemical Ecology 37: 98-104.


Menzel F., Woywod M., Blüthgen N. & Schmitt T. (2010) Behavioural and chemical mechanisms behind a Mediterranean ant-ant association. Ecological Entomology 35: 711-720.

Drescher J., Blüthgen N., Schmitt T., Bühler J. & Feldhaar H. (2010) Societies drifting apart? Behavioral, genetic and chemical differentiation between supercolonies in the yellow crazy ant Anaplolepis gracilipesPlosOne 5: e13581.

Leonhardt S., Zeilhofer S., Blüthgen N. & Schmitt T. (2010) Singless bees use terpenes as olfactory cues to find resin sources. Chemical Senses 35: 593-601.

Leonhardt S., Jung L.-M., Schmitt T. & Blüthgen N. (2010) Terpenoids tame aggressors: role of chemicals in stingless bee communal nesting. Behavioral Ecology & Sociobiology 64: 1415-1523.

Menzel F., Pokorny T., Blüthgen N. & Schmitt T. (2010) Trail-sharing among tropical ants: interspecific use of trail pheromones? Ecological Entomology 35: 495-503.

Kaltenpoth M., Schmitt T., Polidori C., Koedam D. & Strohm E. (2010) Symbiotic streptomycetes in the antennae of South American Trachypus digger wasps (Hymenoptera, Carbronidae). Physiological Entomology 35: 196-200.

Raychoudhury R., Desjardins C. A., Buellesbach J., Loehlin D. W., Grillenberger B. K., Beukeboom L., Schmitt T. & Werren J. H. (2010) Behavioural and genetic characteristics of a new species of NasoniaHeredity 104: 278-288.

The Nasonia Genome Working Group (2010) Functional and evolutionary insights from the genomes of three Nasonia species. Science327: 343-348.


Kaltenpoth M., Schmitt T. & Strohm E. (2009) Hydrocarbons of the antennal gland secretion of female European beewolves, Philanthus triangulum (Hymenoptera: Crabronidae). Chemoecology 19: 219-225.

Menzel F., Schmitt T. & Blüthgen N. (2009) Interspecific nestmate recognition in two parabiotic ant species: acquired recognition cues and low inter-colony discrimination. Insectes Sociaux 56: 251-260.

Leonhardt S., Blüthgen N. & Schmitt T. (2009) Smelling like resin: terpenoids account for species-specific cuticular profiles in Southeast-Asian stingless bees. Insectes Sociaux 56: 157-170.

Kroiss J., Schmitt T. & Strohm E. (2009) Low level of cuticular hydrocarbons in a parasitoid of a solitary digger wasp and its potential for concealment. Entomological Science 12: 9-16.

Geiselhardt S., Schmitt T. & Peschke K. (2009) Chemical composition and pheromonal function of the defensive secretions in the subtribe Stizopina (Coleoptera, Tenebrionidae, Opatrini). Chemoecology 19: 1-6.


Menzel F., Blüthgen N. & Schmitt T. (2008) Tropical parabiotic ants: Highly unusual cuticular substances and low interspecific discrimination. Frontiers in Zoology 5:16.

Strohm E., Herzner G., Kaltenpoth, M., Boland W., Schreier P., Geiselhardt S., Peschke K. & Schmitt T. (2008) The chemistry of the postpharyngeal gland of female European Beewolves. Journal of Chemical Ecology 34: 575-583.

Oettler J., Schmitt T., Herzner G. & Heinze J. (2008) Chemical profiles of mated and virgin queens, egg-laying intermorphs and workers of the ant Crematogaster smithiiJournal of Insect Physiology 54: 672-679.

Haberer W., Schmitt T., Peschke K., Schreier P. & Müller J. K. (2008) Ethyl 4-methyl heptanoate: a male-produced pheromone of Nicrophorus vespilloidesJournal of Chemical Ecology 34: 94-98.

Strohm E., Kroiss J., Herzner G., Laurien-Kehnen C., Boland W., Schreier P. & Schmitt T. (2008): A cuckoo in wolves’ clothing? Chemical mimicry in a specialized cuckoo wasp of the European beewolf (Hymenoptera, Chrysididae and Crabronidae). Frontiers in Zoology 5: 2.


Herzner G., Schmitt T., Peschke K., Hilpert A. & Strohm E. (2007) Food wrapping with the postpharyngeal gland secretion by females of the European beewolf Philanthus triangulumJournal of Chemical Ecology 33: 849-859.

Schmitt T., Herzner G., Weckerle B., Schreier P. & Strohm E. (2007) Volatiles of foraging honeybees Apis mellifera L. (Hymenoptera: Apidae) and their potential role as semiochemicals. Apidologie 38: 164-170.


Kroiss J., Schmitt T., Schreier P., Strohm E. & Herzner G. (2006) A selfish function of a social gland? A postpharyngeal gland functions as sex pheromone reservoir in males of a solitary wasp. Journal of Chemical Ecology 32: 2763-2776.

Brandt M., Heinze J., Schmitt T. & Foitzik, S. (2006) Convergent evolution of the Dufour’s gland secretion as a propaganda substance in the slave-making ant genera Protomognathus and HarpagoxenusInsectes Sociaux 53: 291-299.

Herzner G., Schmitt T., Schreier P. & Strohm E. (2006) Brothers smell similar: Variation in the sex pheromone of male European Beewolves and its implications for inbreeding avoidance. Biological Journal of the Linnean Society 89: 433-442.


Böll S., Schmitt T., Burschka C., Schreier P., Schwappach P. & Herrmann J. V. (2005): Calciumtartrate crystals in the midgut of the grape leafhopper. Journal of Chemical Ecology 31: 2847-2856.

Herzner G., Schmitt T., Linsenmair K. E. & Strohm E. (2005) Prey recognition by females of the European beewolf and its potential for a sensory trap. Animal Behaviour 70: 1411-1418.

Brandt M., Heinze J., Schmitt T. & Foitzik, S. (2005) A chemical level in the coevolutionary arms race between an ant social parasite and its hosts. Journal of Evolutionary Biology 18: 576-586.


Schmitt T., Krell F.-T. & Linsenmair K. E. (2004) Quinone mixture as attractant for necrophagous dung beetles on dead millipedes. Journal of Chemical Ecology 30: 731-740.

Endler A., Liebig J., Schmitt T., Parker J., Jones G., Schreier P. & Hölldobler B. (2004) Surface hydrocarbons of queen eggs regulate worker reproduction in a social insect. Proceedings of the National Academy of Science of the USA 101: 2945-2950.


Herzner G., Schmitt T., Linsenmair K. E. & Strohm E. (2003) Flagellar sensilla in male and female European beewolves, Philanthus triangulum F. (Hymenoptera: Sphecidae). Entomologica Fennica 14: 237–247.

Schmitt T., Strohm E., Herzner G., Bicchi C., Krammer G., Heckel F. & Schreier P. (2003) (S)-2,3-Dihydrofarnesoic acid, a new component in cephalic glands of male European beewolves Philanthus triangulumJournal of Chemical Ecology 29: 2469-2479.


Schmitt T. & Lehman N. (1999) Non-unity molecular heritability demonstrated by continuous evolution in vitro. Chemistry & Biology 6: 857-869.

Krell F.-T., Schmitt T. & Linsenmair K. E. (1999) The rhinoceros beetle Cyphonistes tuberculifrons Quedenfeldt 1884 attracted by quinones (Coleoptera Scarabaeidae Dynastinae): primary olfactory attraction of a saprophagous beetle by its source of food? Tropical Zoology 12: 297-308.


Krell F.-T. Schmitt T. & Linsenmair K. E. (1997): Diplopod defensive secretions as attractants for necrophagous scarab beetles (Diplopoda; Insecta, Coleoptera: Scarabaeidae). Entomologica Scandinavica Supplement 51: 281-285.