Journal of Ethology

, Volume 28, Issue 2, pp 295–304 | Cite as

Male territoriality and mating system in the European beewolf Philanthus triangulum F. (Hymenoptera: Crabronidae): evidence for a “hotspot” lek polygyny

  • Johannes KroissEmail author
  • Klaus Lechner
  • Erhard Strohm


The evolution of the mating system of a species is strongly influenced by the spatial and temporal distribution of females and/or resources. Here, we describe aspects of the territorial behavior of males of a solitary digger wasp, the European beewolf (Philanthus triangulum) and characterize the mating system. We show that beewolf males establish small territories that do not contain any resources essential to females. These territories are intensively scent-marked with a pheromone from a cephalic gland and are defended against intruders in combat flights. We provide evidence that scent-marking constitutes a chemical display and that the pheromone serves to attract receptive females and, thus, represents a sex pheromone. Using spatial statistics, we show that beewolf territories are clumped in space both with respect to other male territories and, more importantly, with respect to female nesting sites. Additionally, the proportion of days a territory is occupied by a male is correlated with the number of female nests in the vicinity. Taking into account that beewolf males do not defend or provide resources essential to females, but merely display chemically to attract females for mating in an aggregation of territories close to female nesting sites, we conclude that the European beewolf exhibits a hotspot lek polygyny with female nesting sites constituting “hotspots” for lek formation.


Scent-marking Sex pheromone Sphecidae Digger wasp 



This study was partly supported by the DFG (STR 532/1-2). We thank Thorsten Wiegand for providing the Programita Software. We are grateful to anonymous reviewers who provided valuable comments on an earlier draft of the manuscript.


  1. Alcock J (1974) The behaviour of Philanthus crabroniformis (Hymenoptera: Sphecidae). J Zool 33:173–246Google Scholar
  2. Alcock J (1975a) Male mating strategies of some philanthine wasps (Hymenoptera: Sphecidae). J Kansas Entomol Soc 48:532–545Google Scholar
  3. Alcock J (1975b) Territorial behavior by males of Philanthus multimaculatus (Hymenoptera-Sphecidae) with a review of territoriality in male sphecids. Anim Behav 23:889–895CrossRefGoogle Scholar
  4. Alcock J (1981) Lek territoriality in the tarantula hawk wasp Hemipepsis ustulata (Hymenoptera, Pompilidae). Behav Ecol Sociobiol 8:309–317CrossRefGoogle Scholar
  5. Alcock J, Bailey W (1997) Success in territorial defence by male tarantula hawk wasps Hemipepsis ustulata: the role of residency. Ecol Entomol 22:377–383CrossRefGoogle Scholar
  6. Alcock J, Carey M (1988) Hilltopping behavior and mating success of the tarantula hawk wasp, Hemipepsis ustulata (Hymenoptera, Pompilidae), at a high elevation peak. J Nat Hist 22:1173–1178CrossRefGoogle Scholar
  7. Alcock J, Johnson MD (1990) Female mate choice in the carpenter bee Xylocopa varipuncta (Hymenoptera: Anthophoridae). J Zool 221:195–204CrossRefGoogle Scholar
  8. Alcock J, Kemp DJ (2006) The behavioral significance of male body size in the tarantula hawk wasp Hemipepsis ustulata (Hymenoptera: Pompilidae). Ethol 112:691–698CrossRefGoogle Scholar
  9. Alcock J, Jones CE, Buchmann SL (1977) Male mating strategies in the bee Centris pallida Fox (Anthophoridae-Hymenoptera). Am Nat 111:145–155CrossRefGoogle Scholar
  10. Baker RR (1983) Insect territoriality. Annu Rev Entomol 28:65–89CrossRefGoogle Scholar
  11. Beani L, Sledge MF, Maiani S, Boscaro F, Landi M, Fortunato A, Turillazzi S (2002) Behavioral and chemical analyses of scent-marking in the lek system of a hover-wasp (Vespidae, Stenogastrinae). Insectes Soc 49:275–281CrossRefGoogle Scholar
  12. Birkhofer K, Henschel JR, Scheu S (2006) Spatial-pattern analysis in a territorial spider: evidence for multi-scale effects. Ecography 29:641–648CrossRefGoogle Scholar
  13. Bradbury JW (1977) Lek mating-behavior in hammer-headed bat. Z Tierpsychol 45:225–255Google Scholar
  14. Bradbury JW (1981) The evolution of leks. In: Alexander RD, Tinkle DW (eds) Natural selection and social behavior. Chiron Press, New York, pp 138–169Google Scholar
  15. Brown JL, Orians GH (1970) Spacing patterns in mobile animals. Annu Rev Ecol Syst 1:239–262CrossRefGoogle Scholar
  16. Coelho JR, Holliday CW (2001) Effects of size and flight performance on intermale mate competition in the cicada killer, Sphecius speciosus Drury (Hymenoptera: Sphecidae). J Insect Behav 14:345–351CrossRefGoogle Scholar
  17. Diggle P (1979) On parameter estimation and goodness-of-fit testing for spatial point patterns. Biometrics 35:87–101CrossRefGoogle Scholar
  18. Emlen ST, Oring LW (1977) Ecology, sexual selection, and the evolution of mating systems. Science 197:215–223CrossRefPubMedGoogle Scholar
  19. Evans HE (1982) Nesting and territorial behavior of Philanthus barbatus Smith (Hymenoptera, Sphecidae). J Kansas Entomol Soc 55:571–576Google Scholar
  20. Evans HE, O’Neill KM (1978) Alternative mating strategies in digger wasp Philanthus zebratus Cresson. PNAS 75:1901–1903CrossRefPubMedGoogle Scholar
  21. Evans HE, O’Neill KM (1988) The natural history of North American beewolves. Cornell University Press, IthacaGoogle Scholar
  22. Field S, Kaspi R, Yuval B (2002) Why do dalling medflies (Diptera: Tephritidae) cluster? Assessing the empirical evidence for models of medfly lek evolution. Fla Entomol 85:63–72CrossRefGoogle Scholar
  23. Goettler W, Strohm E (2008) Mandibular glands of European beewolves, Philanthus triangulum (Hymenoptera, Crabronidae). Arth Struct Dev 37:363–371CrossRefGoogle Scholar
  24. Gwynne DT (1978) Male territoriality in the bumble bee wolf Philanthus bicinctus (Mickel) (Hymenoptera: Sphecidae): observations on the behaviour of individual males. Z Tierpsychol 47:89–103Google Scholar
  25. Gwynne DT (1980) Female defense polygyny in the bumblebee wolf, Philanthus bicinctus (Hymenoptera, Sphecidae). Behav Ecol Sociobiol 7:213–225CrossRefGoogle Scholar
  26. Herzner G, Schmitt T, Linsenmair KE, Strohm E (2005) Prey recognition by females of the European beewolf and its potential for a sensory trap. Anim Behav 70:1411–1418CrossRefGoogle Scholar
  27. Herzner G, Schmitt T, Heckel F, Schreier P, Strohm E (2006) Brothers smell similar: variation in the sex pheromone of male European beewolves Philanthus triangulum F. (Hymenoptera: Crabronidae) and its implications for inbreeding avoidance. Biol J Linnean Soc 89:433–442CrossRefGoogle Scholar
  28. Höglund J, Alatalo RV (1995) Leks. Princeton University Press, PrincetonGoogle Scholar
  29. Hovi M, Alatalo RV, Halonen M, Lundberg A (1997) Responses of male and female black grouse to male vocal display. Ethology 103:1032–1041CrossRefGoogle Scholar
  30. Kaltenpoth M, Strohm E (2006) The scent of senescence: age-dependent changes in the composition of the marking pheromone of the male European beewolf, Philanthus triangulum. J Insect Sci 6:20CrossRefGoogle Scholar
  31. Kaltenpoth M, Kroiss J, Strohm E (2007) The odor of origin: kinship and geographical distance are reflected in the marking pheromone of male beewolves (Philanthus triangulum, Hymenoptera, Crabronidae). BMC Ecol 7:11CrossRefPubMedGoogle Scholar
  32. Kaspi R, Yuval B (1999) Lek site selection by male Mediterranean fruit flies. J Insect Behav 12:267–276CrossRefGoogle Scholar
  33. Kemp DJ, Alcock J (2003) Lifetime resource utilization, flight physiology, and the evolution of contest competition in territorial insects. Am Nat 162:290–301CrossRefPubMedGoogle Scholar
  34. Kemp DJ, Krockenberger AK (2004) Behavioural thermoregulation in butterflies: the interacting effects of body size and basking posture in Hypolimnas bolina (L.) (Lepidoptera: Nymphalidae). Aust J Zool 52:229–236CrossRefGoogle Scholar
  35. Kemp D, Wiklund C (2001) Fighting without weaponry: a review of male-male contest competition in butterflies. Behav Ecol Sociobiol 49:429–442CrossRefGoogle Scholar
  36. Kemp DJ, Alcock J, Allen GR (2006a) Sequential size assessment and multicomponent decision rules mediate aerial wasp contests. Anim Behav 71:279–287CrossRefGoogle Scholar
  37. Kemp DJ, Wiklund C, Gotthard K (2006b) Life history effects upon contest behaviour: age as a predictor of territorial contest dynamics in two populations of the speckled wood butterfly, Pararge aegeria L. Ethology 112:471–477CrossRefGoogle Scholar
  38. Kemp DJ, Wiklund C, van Dyck H (2006c) Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance. Behav Ecol Sociobiol 59:403–411CrossRefGoogle Scholar
  39. Kennedy JS (1983) Zigzagging and casting as a programmed response to wind-borne odour: a review. Physiol Entomol 8:109–120CrossRefGoogle Scholar
  40. Kirkpatrick M, Ryan M (1991) The evolution of mating preferences and the paradox of the lek. Nature 350:33–38CrossRefGoogle Scholar
  41. 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. J Chem Ecol 32:2763–2776CrossRefPubMedGoogle Scholar
  42. Marden J, Rollins R (1994) Assessment of energy reserves by damselflies engaged in aerial contests for mating territories. Anim Behav 48:1023–1030CrossRefGoogle Scholar
  43. Marden J, Waage J (1990) Escalated damselfly territorial contests are energetic wars of attrition. Anim Behav 39:954–959CrossRefGoogle Scholar
  44. Minckley RL, Buchmann SL, Wcislo WT (1991) Bioassay evidence for a sex attractant pheromone in the large carpenter bee, Xylocopa varipuncta (Anthophoridae, Hymenoptera). J Zool 224:285–291CrossRefGoogle Scholar
  45. O’Neill KM (1979) Territorial behavior in males of Philanthus psyche (Hymenoptera: Sphecidae). Psyche 86:19–43CrossRefGoogle Scholar
  46. O’Neill K (2001) Solitary wasps. Behavior and natural history. Comstock Publishing, IthacaGoogle Scholar
  47. O′Neill KM (1983) Territoriality, body size, and spacing in males of the beewolf Philanthus basilaris (Hymenoptera; Sphecidae). Behaviour 86:295–321CrossRefGoogle Scholar
  48. Partecke J, von Haeseler A, Wikelski M (2002) Territory establishment in lekking marine iguanas, Amblyrhynchus cristatus: support for the hotshot mechanism. Behav Ecol Sociobiol 51:579–587CrossRefGoogle Scholar
  49. Raffa KF (2001) Mixed messages across multiple trophic levels: the ecology of bark beetle chemical communication systems. Chemoecology 11:49–65CrossRefGoogle Scholar
  50. Ripley B (1981) Spatial statistics. Wiley, New YorkCrossRefGoogle Scholar
  51. Schroeder MA, White GC (1993) Dispersion of greater prairie chicken nests in relation to lek location: evaluation of the hot-spot hypothesis of lek evolution. Behav Ecol 4:266–270CrossRefGoogle Scholar
  52. Shelly T, Whittier T (1997) Lek behavior of insects. In: Choe J, Crespi B (eds) Mating systems in insects and arachnids. Cambridge University Press, Cambridge, pp 273–293CrossRefGoogle Scholar
  53. Shorey HH (1973) Behavioral responses to insect pheromones. Annu Rev Entomol 18:349–380CrossRefPubMedGoogle Scholar
  54. Simon-Thomas RT, Poorter EPR (1972) Notes on the behavior of males of Philanthus triangulum (Hymenoptera, Sphecidae). Tijdschr Entomol 115:141–151Google Scholar
  55. Stoyan D, Stoyan H (1994) Fractals, random shapes and point fields. Methods of geometrical statistics. Wiley, New YorkGoogle Scholar
  56. Strohm E, Lechner K (2000) Male size does not affect territorial behaviour and life history traits in a sphecid wasp. Anim Behav 59:183–191CrossRefPubMedGoogle Scholar
  57. Strohm E, Linsenmair KE (1997) Low resource availability causes extremely male-biased investment ratios in the European beewolf, Philanthus triangulum F. (Hymenoptera, Sphecidae). Proc R Soc Lond B 264:423–429CrossRefGoogle Scholar
  58. Strohm E, Linsenmair KE (1999) Measurement of parental investment and sex allocation in the European beewolf Philanthus triangulum F. (Hymenoptera: Sphecidae). Behav Ecol Sociobiol 47:76–88CrossRefGoogle Scholar
  59. Strohm E, Marliani A (2002) The cost of parental care: prey hunting in a digger wasp. Behav Ecol 13:52–58CrossRefGoogle Scholar
  60. Takeuchi T (2006) Matter of size or matter of residency experience? Territorial contest in a green hairstreak, Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae). Ethology 112:293–299CrossRefGoogle Scholar
  61. Velthuis HHW, De Camargo JMF (1974) Further observations on the function of male territories in the carpenter bee Xylocopa (Neoxylocopa) hirsutissima Maidl (Anthophoridae, Hymenoptera). Neth J Zool 25:516–528CrossRefGoogle Scholar
  62. Velthuis HHW, Gerling D (1980) Observations on territoriality and mating behaviour of the carpenter bee Xylocopa sulcatipes. Entomol Exp Appl 28:82–91CrossRefGoogle Scholar
  63. Vinson SB, Frankie GW (1990) Territorial and mating-behavior of Xylocopa fimbriata F and Xylocopa gualanensis Cockerell from Costa-Rica. J Insect Behav 3:13–32CrossRefGoogle Scholar
  64. Weldon C (2007) Influence of male aggregation size on female visitation in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae). Aust J Entomol 46:29–34CrossRefGoogle Scholar
  65. Westcott D (1994) Leks of leks: a role for hotspots in lek evolution? Proc R Soc Lond B 258:281–286CrossRefGoogle Scholar
  66. Wiegand T, Moloney K (2004) Rings, circles, and null-models for point pattern analysis in ecology. Oikos 104:209–229CrossRefGoogle Scholar
  67. Wiegand T, Jeltsch F, Ward D (2000) Do spatial effects play a role in the spatial distribution of desert-dwelling Acacia raddiana? J Veg Sci 11:473–484CrossRefGoogle Scholar
  68. Wiley R (1974) Evolution of social organization and life history patterns among grouse (Aves: Tetraonidae). Q Rev Biol 49:201–227CrossRefPubMedGoogle Scholar
  69. Young K, Genner M, Joyce D, Haesler M (2009) Hotshots, hot spots, and female preference: exploring lek formation models with a bower-building cichlid fish. Behav Ecol 20:609–615CrossRefGoogle Scholar

Copyright information

© Japan Ethological Society and Springer 2009

Authors and Affiliations

  1. 1.Institute of ZoologyUniversity of RegensburgRegensburgGermany
  2. 2.Department of Animal Ecology and Tropical BiologyUniversity of WürzburgWürzburgGermany

Personalised recommendations