Effects of Pheromone Dose and Conspecific Density on the Use of Aggregation-Sex Pheromones by the Longhorn Beetle Phymatodes grandis and Sympatric Species (Coleoptera: Cerambycidae)

  • R. Maxwell CollignonEmail author
  • Jonathan A. Cale
  • J. Steven McElfresh
  • Jocelyn G. Millar


Many species of longhorn beetles (Coleoptera: Cerambycidae) utilize male-produced aggregation-sex pheromones that attract both sexes. However, the reasons why and the details of how this type of pheromone is used by cerambycids and other coleopteran species that utilize analogous male-produced pheromones remain unclear. Thus, our goals were to test the hypotheses that 1) cerambycids respond to pheromones in a dose-dependent (= release rate-dependent) manner and 2) pheromone emission is density-dependent. If true, these characteristics of pheromone use could suggest that cerambycids utilize an optimal density strategy to limit competition for scarce and ephemeral hosts, i.e., the stressed or dying trees that typically constitute their larval hosts. Attraction of beetles to a range of release rates of two common pheromone components – 2-methylbutanol and 3-hydroxyhexan-2-one – was tested in field trials. Responses, as measured by the number of beetles caught in pheromone-baited traps, increased with release rates for five endemic species, even at the highest rates tested (~1450 μg/h for 2-methylbutanol and ~720 μg/h for 3-hydroxyhexan-2-one). The effect of density of conspecific males on per capita pheromone production was tested by collecting the volatiles produced by individuals, pairs, or groups of three or four male Phymatodes grandis Casey. Frequency of pheromone production was significantly different among the treatment densities, and emission rates of the pheromone (R)-2-methylbutanol decreased with increasing density. These results are discussed in the context of a possible optimal density strategy used by cerambycids, and more broadly, in relation to the use of male-produced aggregation-sex pheromones by other coleopterans. In addition, we report the identification of the pheromones of four of our five test species, specifically, Phymatodes obliquus Casey, Brothylus conspersus LeConte, Brothylus gemmulatus LeConte, and Xylotrechus albonotatus Casey.


Cerambycidae Aggregation pheromone Male-produced pheromone Density effects 



We thank Jacqueline Serrano for assistance with fieldwork, and Ian Swift for consultation regarding cerambycid biology. We also thank Dr. Lawrence Hanks for feedback and comments on the manuscript, the anonymous reviewers of the submitted manuscript for their constructive criticism, and Dr. David Hall for his thoughtful edits. We gratefully acknowledge the financial support for this research provided by the USDA-AFRI-NIFA Predoctoral Fellowship Program to RMC (2016-67011-25097).

Supplementary material

10886_2019_1047_MOESM1_ESM.docx (483 kb)
ESM 1 (DOCX 483 kb)


  1. Arnaud L, Lognay G, Verscheure M, Leenaers L, Gaspar C, Haubruge E (2002) Is dimethyldecanal a common aggregation pheromone of Tribolium flour beetles? J Chem Ecol 28:523–532CrossRefGoogle Scholar
  2. Baker TC, Cardé RT, Miller JR (1980) Oriental fruit moth pheromone component emission rates measured after collection by glass-surface adsorption. J Chem Ecol 6:749–758CrossRefGoogle Scholar
  3. Bartelt RJ, James DG (1994) Aggregation pheromone of Australian sap beetle, Carpophilus davidsoni (Coleoptera: Nitidulidae). J Chem Ecol 20:3207–3219CrossRefGoogle Scholar
  4. Bartelt RJ, Weaver DK, Arbogast RT (1995) Aggregation pheromone of Carpophilus dimidiatus (F.) (Coleoptera: Nitidulidae) and responses to Carpophilus pheromones in South Carolina. J Chem Ecol 21:1763–1779CrossRefGoogle Scholar
  5. Bengtsson J (2008) Aggregation in non-social insects. Växtskyddsbiologi, Sveriges lantbruksuniversitet. Alnarp, Sweden, pp 1–18Google Scholar
  6. Byers JA (1983) Sex-specific responses to aggregation pheromone: regulation of colonization density in the bark beetle Ips paraconfusus. J Chem Ecol 9:129–142CrossRefGoogle Scholar
  7. Cardé RT (2014) Defining attraction and aggregation pheromones: teleological versus functional perspectives. J Chem Ecol 40:519–520CrossRefGoogle Scholar
  8. Collignon RM, Swift IP, Zou Y, McElfresh JS, Hanks LM, Millar JG (2016) The influence of host plant volatiles on the attraction of longhorn beetles to pheromones. J Chem Ecol 42:215–229CrossRefGoogle Scholar
  9. Goldsmith SK, Stewart Z, Adams S, Trimble A (1996) Body size, male aggression, and male mating success in the cottonwood borer, Plectrodera scalator (Coleoptera: Cerambycidae). J Insect Behav 9:719–727CrossRefGoogle Scholar
  10. Graham EE, Mitchell RF, Reagel PF, Barbour JD, Millar JG, Hanks LM (2010) Treating panel traps with a fluoropolymer enhances their efficiency in capturing cerambycid beetles. J Econ Entomol 103:641–647CrossRefGoogle Scholar
  11. Hanks LM, Millar JG (2016) Sex and aggregation-sex pheromones of cerambycid beetles: basic science and practical applications. J Chem Ecol 42:631–654CrossRefGoogle Scholar
  12. Hanks LM, Millar JG, Moreira JA, Barbour JD, Lacey ES, McElfresh JS, Reuter FR, Ray AM (2007) Using generic pheromone lures to expedite identification of aggregation pheromones for the cerambycid beetles Xylotrechus nauticus, Phymatodes lecontei, and Neoclytus modestus modestus. J Chem Ecol 33:889–907CrossRefGoogle Scholar
  13. Hanks LM, Millar JG, Mongold-Diers JA, Wong JCH, Meier LR, Reagel PF, Mitchell RF (2012) Using blends of cerambycid beetle pheromones and host plant volatiles to simultaneously attract a diversity of cerambycid species. Can J For Res 42:1050–1059CrossRefGoogle Scholar
  14. Hayes RA, Griffiths MW, Nahrung HF, Arnold PA, Hanks LM, Millar JG (2016) Optimizing generic cerambycid pheromone lures for Australian biosecurity and biodiversity monitoring. J Econ Entomol 109:1741–1749CrossRefGoogle Scholar
  15. Hock V, Chouinard G, Lucas É, Cormier D, Leskey T, Wright S, Zhang A, Pichette A (2014) Establishing abiotic and biotic factors necessary for reliable male pheromone production and attraction to pheromones by female plum curculios Conotrachelus nenuphar (Coleoptera: Curculionidae). Can Entomol 146:528–547CrossRefGoogle Scholar
  16. Imrei Z, Millar JG, Janik G, Tóth M (2013) Field screening of known pheromone components of longhorned beetles in the subfamily Cerambycinae (Coleoptera: Cerambycidae) in Hungary. Zeitschrift fur Naturforsch - Sect C J Biosci 68(C):236–242CrossRefGoogle Scholar
  17. Jones TM, Quinnell R (2002) Testing predictions for the evolution of lekking in the sandfly, Lutzomyia longipalpis. Anim Behav 63:605–612CrossRefGoogle Scholar
  18. Lacey ES, Ginzel MD, Millar JG, Hanks LM (2004) Male-produced aggregation pheromone of the cerambycid beetle Neoclytus acuminatus acuminatus. J Chem Ecol 30:1493–1507CrossRefGoogle Scholar
  19. Lacey ES, Moreira JA, Millar JG, Ray AM, Hanks LM (2007) Male-produced aggregation pheromone of the cerambycid beetle Neoclytus mucronatus mucronatus. Entomol Exp Appl 122:171–179CrossRefGoogle Scholar
  20. Landolt PJ (1997) Sex attractant and aggregation pheromones of male phytophagous insects. Am Entomol 42:12–22CrossRefGoogle Scholar
  21. Lemay MA, Silk PJ, Sweeney J (2010) Calling behavior of Tetropium fuscum (Coleoptera: Cerambycidae: Spondylidinae). Can Entomol 142:256–260CrossRefGoogle Scholar
  22. Linsley EG, Chemsak JA (1997) The Cerambycidae of North America, part VIII: bibliography, index, and host plant index. University of California Press, BerkeleyGoogle Scholar
  23. Mitchell RF, Graham EE, Wong JCH, Reagel PF, Striman BL, Hughes GP, Paschen MA, Ginzel MD, Millar JG, Hanks LM (2011) Fuscumol and fuscumol acetate are general attractants for many species of cerambycid beetles in the subfamily Lamiinae. Entomol Exp Appl 141:71–77CrossRefGoogle Scholar
  24. Pellegrino AC, Peñaflor MFGV, Nardi C, Bezner-Kerr W, Guglielmo CG, Bento JMS, McNeil JN (2013) Weather forecasting by insects: modified sexual behaviour in response to atmospheric pressure changes. PLoS One 8:e75004. CrossRefGoogle Scholar
  25. Petroski RJ, Bartelt RJ, Vetter RS (1994) Male-produced aggregation pheromone of Carpophilus obseltus (Coleoptera: Nitidulidae). J Chem Ecol 20:1483–1493CrossRefGoogle Scholar
  26. Raffa KF (2001) Mixed messages across multiple trophic levels: the ecology of bark beetle chemical communication systems. Chemoecology 11:49–65CrossRefGoogle Scholar
  27. Ray AM, Millar JG, Moreira JA, McElfresh JS, Mitchell RF, Barbour JD, Hanks LM (2015) North American species of cerambycid beetles in the genus Neoclytus share a common hydroxyhexanone-hexanediol pheromone structural motif. J Econ Entomol 108:1860–1868CrossRefGoogle Scholar
  28. Sanders CJ (1997) Mechanisms of mating disruption in moths. In: Cardé RT, Minks AK (eds) Insect pheromone research. Springer, Boston, MA, pp 333–346CrossRefGoogle Scholar
  29. Schlyter F, Byers JA, Löfqvist J (1987) Attraction to pheromone sources of different quantity, quality, and spacing: density-regulation mechanisms in bark beetle Ips typographus. J Chem Ecol 13(6):1503–1523CrossRefGoogle Scholar
  30. Scholz D, Tchabi A, Markham RH, Poehling HM, Borgemeister C (1998) Factors affecting pheromone production and behavioral responses by Prostephanus truncatus (Coleoptera: Bostrichidae). Ann Entomol Soc Am 91:872–878CrossRefGoogle Scholar
  31. Smith JL, Cork A, Hall DR, Hodges RJ (1996) Investigation of the effect of female larger grain borer, Prostephanus truncates (horn) (Coleoptera: Bostrichidae), and their residues on the production of aggregation pheromone by males. J Stored Prod Res 32:171–181CrossRefGoogle Scholar
  32. Sweeney JD, Silk PJ, Gutowski JM, Wu J, Lemay MA, Mayo PD, Magee DI (2010) Effect of chirality, release rate, and host volatiles on response of Tetropium fuscum (F.), Tetropium cinnamopterum Kirby, and Tetropium castaneum (L.) to the aggregation pheromone, fuscumol. J Chem Ecol 36:1309–1321CrossRefGoogle Scholar
  33. The R Foundation (2016) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria Google Scholar
  34. Venables WN, Ripley BD (2002) Modern applied statistics with S, Fourth edn. Springer, New YorkGoogle Scholar
  35. Weatherston I, Miller D, Lavoie-Dornik J (1985) Capillaries as controlled release devices for insect pheromones and other volatile substances - a reevaluation: part II. Predicting release rates from Celcon and Teflon capillaries. J Chem Ecol 11:967–978CrossRefGoogle Scholar
  36. Wertheim B, van Baalen EJA, Dicke M, Vet LEM (2005) Pheromone-mediated aggregation in nonsocial arthropods: an evolutionary perspective. Annu Rev Entomol 50:321–346CrossRefGoogle Scholar
  37. Wickham JD, Harrison RD, Lu W, Guo Z, Millar JG, Hanks LM, Chen Y (2014) Generic lures attract cerambycid beetles in a tropical montane rain forest in southern China. J Econ Entomol 107:259–267CrossRefGoogle Scholar
  38. Williams RN, Ellis MS, Bartelt RJ (1995) Efficacy of Carpophilus aggregation pheromones on nine species in northeastern Ohio, and identification of the pheromone of C. brachypterus. Entomol Exp Appl 77:141–147CrossRefGoogle Scholar
  39. Wong JCH, Meier LR, Zou Y, Mongold-Diers JA, Hanks LM (2017) Evaluation of Methods Used in Testing Attraction of Cerambycid Beetles to Pheromone-Baited Traps. J Econ Entomol 110:2269–2274Google Scholar
  40. Wood DL (1982) The role of pheromones, kairomones, and allomones in the host selection and colonization behavior of bark beetles. Annu Rev Entomol 27:411–446CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Department of EntomologyUniversity of CaliforniaRiversideUSA
  2. 2.USDA-ARS, Pacific Basin Ag. Res. CenterHiloUSA
  3. 3.Department of Renewable ResourcesUniversity of AlbertaEdmontonCanada

Personalised recommendations