Journal of Insect Behavior

, Volume 26, Issue 3, pp 304–320 | Cite as

Mating Behavior and Performance in the Two Cosmopolitan Mealybug Species Planococcus citri and Pseudococcus calceolariae (Hemiptera: Pseudococcidae)

  • Elsa Borges da Silva
  • Manuela Branco
  • Zvi Mendel
  • José Carlos Franco


Mating strategies of mealybugs were investigated using two heterogeneric cosmopolitan species as case study: Planococcus citri (Risso) and Pseudococcus calceolariae (Maskell). Male mating behavior of the studied species differed in respect to mate selection, and frequency and duration of copulation. Females played an active role in mate selection, by either facilitating or resisting copulation. This is a first evidence of female sexual active behavior in scale insects. In both species, male predisposition to mate was affected by previous exposition to light. The frequency of courtship and copulation decreased with the age of females for Pl. citri but not for Ps. calceolariae. Mating frequency increased with female colony density in both species.


Coccoidea courtship mating system female choice Planococcus citri Pseudococcus calceolariae 



We thank Manuel Cariano and Leonor Campos for insect rearing and laboratory assistance. We also acknowledge the comments and suggestions of two anonymous reviewers on an earlier version of the manuscript. This research was supported by the Fundação para a Ciência e Tecnologia and FEDER (projects POCI/AGR/57580/2004, and PTDC/AGR-AAM/099560/2008; the first author received two research grants, SFRH/BPD/22145/2005, and SFRH/BPD/44923/2008).


  1. Andersson M, Simmons LW (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296–302PubMedCrossRefGoogle Scholar
  2. Arnaud L (1999) La compétition spermatique chez les insectes: les stratégies d’assurance de la paternité et la préséance du sperme. Biotechnol Agron Soc Environ 3:86–103Google Scholar
  3. Arnqvist G, Nilsson T (2000) The evolution of polyandry: multiple mating and female fitness in insects. Anim Behav 60:145–164PubMedCrossRefGoogle Scholar
  4. Ben-Dov Y (1994) A systematic catalogue of the mealybugs of the world (Insecta: Homoptera: Coccoidea: Pseudoccocidae and Putoidae) with data on their geographical distribution, host plants, biology and economic importance. Intercept, AndoverGoogle Scholar
  5. Bierl-Leonhardt BA, Moreno DS, Schwarz M, Fargerlund J, Plimmer JR (1981) Isolation, identification and synthesis of the sex-pheromone of the citrus mealybug, Planococcus citri (Risso). Tetrahedron Lett 22:389–392CrossRefGoogle Scholar
  6. Boake CRB, Shelly TE, Kaneshiro KY (1996) Sexual selection in relation to pest-management strategies. Annu Rev Entomol 41:211–229PubMedCrossRefGoogle Scholar
  7. Branco M, Jactel H, Franco JC, Mendel Z (2006) Modelling response to insect trap captures to pheromone dose. Ecol Model 197:247–257CrossRefGoogle Scholar
  8. Cabaleiro C, Segura A (1997) Field transmission of grapevine leafroll associated virus 3 (GLRaV-3) by the mealybug Planococcus citri. Plant Dis 81:283–287CrossRefGoogle Scholar
  9. Chong JH, Roda AL, Mannion CM (2008) Life history of the mealybug, Maconellicoccus hirsutus (Hemiptera: Pseudococcidae), at constant temperatures. Environ Entomol 37:323–332PubMedCrossRefGoogle Scholar
  10. El-Minshawy AM, Karan HH, El-Sawaf SK (1974) Biological studies on the long tailed meaybug Pseudococcus longispinus (Targ. and Tozzeti). Bull Entomol Soc Egypt 58:385–391Google Scholar
  11. El-Sayed AM (2011) The pherobase: database of insect pheromones and semiochemicals. <>
  12. El-Sayed AM, Unelius CR, Twidle A, Mitchell V, Manning LA, Cole L, Suckling DM, Flores MF, Zaviezo T, Bergmann J (2010) Chrysanthemyl 2-acetoxy-3-methylbutanoate: the sex pheromone of the citrophilous mealybug. Pseudococcus calceolariae. Tetrahedron Lett 51:1075–1078CrossRefGoogle Scholar
  13. Emlen ST, Oring LW (1977) Ecology, sexual selection, and the evolution of mating systems. Science 197:215–223PubMedCrossRefGoogle Scholar
  14. Franco JC, Marotta S (1999) A survey of the mealybugs in citrus groves in Portugal. Entomologica 33:191–196Google Scholar
  15. Franco JC, Suma P, Silva EB, Blumberg D, Mendel Z (2004) Management strategies of mealybug pests of citrus in Mediterranean countries. Phytoparasitica 32:507–522CrossRefGoogle Scholar
  16. Franco JC, Zada A, Mendel Z (2009) Novel approaches for the management of mealybug pests. In: Ishaaya I, Horowitz AR (eds) Biorational control of Arthropod Pests: application and resistance management. Springer Netherlands, Dordrecht, pp 233–278CrossRefGoogle Scholar
  17. Gavrilets S, Arnqvist G, Friberg U (2001) The evolution of female mate choice by sexual conflict. Proc Roy Soc Lond B Biol Sci 268:531–539CrossRefGoogle Scholar
  18. Gullan PJ, Kosztarab M (1997) Adaptations in scale insects. Ann Rev Entomol 42:23–50CrossRefGoogle Scholar
  19. Hales DF (2005) Physiological determinants of male mating performance in aphids (Hemiptera: Aphididae). Eur J Entomol 102:641–646Google Scholar
  20. Henry LM, Roitber BD, Gillespie DR (2006) Covariance of phenotypically plastic traits induces an adaptive shift in host selection behaviour. Proc R Soc B 273:2893–2899PubMedCrossRefGoogle Scholar
  21. James HC (1937) Sex ratios and the status of the male in pseudococcinae (Hem, Coccidae). Bull Entomol Res 28:429–461CrossRefGoogle Scholar
  22. Lanier GN (1990) Principles of attraction-annihilation: mass trapping and other means. In: Ridgway RL, Silverstein RM, Inscoe MN (eds) Behavior-modifying chemicals for insect management. Applications of pheromones and other attractants. Marcel Dekker Inc, New York, pp 25–45Google Scholar
  23. Mathenge CW, Holford P, Hoffmann JH, Spooner-Hart R, Beattie GAC, Zimmermann HG (2009) The biology of Dactylopius tomentosus (Hemiptera: Dactylopiidae). Bull Entomol Res 99:551–559PubMedCrossRefGoogle Scholar
  24. Mendel Z, Protasov A, Jasrotia P, Silva EB, Levi-Zada A, Franco JC (2012) Sexual maturation and aging of adult male mealybugs (Hemiptera; Pseudococcidae). Bull Entomol Res. doi: 10.1017/S0007485311000605
  25. Mendel Z, Protasov A, Zada A, Assael F, Jasrotia P, Franco JC (2008) Longevity and sexual maturity of an adult male mealybug. In: Branco M, Franco JC, Hodgson CJ (eds) Proceedings of the XI international symposium on scale insect studies, Oeiras (Portugal), 24–27 September 2007. ISA Press, Lisboa, p 231Google Scholar
  26. Millar JG, Daane KM, McElfresh JS, Moreira J, Bentley WJ (2005) Chemistry and applications of mealybug sex pheromones. In: Petroski RJ, Tellez MR, Behle RW (eds) Semiochemicals in pest and weed control. American Chemical Society, Washington DC, pp 11–27CrossRefGoogle Scholar
  27. Moreno DS, Fargerlund J, Ewart WH (1984) Citrus mealybug (Homoptera: Pseudococcidae): behaviour of males in response to sex pheromone in laboratory and field. Ann Entomol Soc Am 77:32–38Google Scholar
  28. Nelson-Rees WA (1959) Triple coitus in the Mealy Bug, Planococcus citri (Risso). Nature 183:479PubMedCrossRefGoogle Scholar
  29. Nelson-Rees WA (1960) A study of sex predetermination in the mealybug Planococcus citri (Risso). J Exp Zool 144:111–137PubMedCrossRefGoogle Scholar
  30. Nestel D, Cohen H, Saphir N, Klein M, Mendel Z (1995) Spatial distribution of scale insects—comparative study using Taylor’s power-law. Environ Entomol 24:506–512Google Scholar
  31. Oku K, Yasuda T (2010) Effects of age and mating on female sex attractant pheromone levels in the sorghum plant bug, Stenotus rubrovittatus (Matsumura). Behav Ecol Sociobiol 64:607–615CrossRefGoogle Scholar
  32. Petersen CL, Charles JG (1997) Transmission of grapevine leafroll-associated closteroviruses by Pseudococcus longispinus and P. calceolariae. Plant Pathol 46:509–515CrossRefGoogle Scholar
  33. Rotundo G, Tremblay E (1976) Observations on the flight of the male of Pseudococcus calceolariae (Mask.) (Homoptera, Coccoidea). Boll Lab Entomol Agrar “Filippo Silvestri” 33:108–112, in ItalianGoogle Scholar
  34. Rotundo G, Tremblay E (1980) Evaluation of the daily rate of sex pheromone release by the females of two mealybug species (Homoptera, Coccoidea, Pseudococcidae). Boll Lab Entomol Agrar “Filippo Silvestri” 37:167–170Google Scholar
  35. Rotundo G, Tremblay E (1981) Scent trailing by virgin females of Pseudococcus calceolariae (Homoptera, Coccoidea, Pseudococcidae). J Chem Ecol 7:85–88CrossRefGoogle Scholar
  36. Serrano MS, Lapointe SL (2002) Evaluation of host plants and a meridic diet for rearing Maconellicoccus hirsutus (Hemiptera: Pseudococcidae) and its parasitoid Anagyrus kamali (Hymenoptera: Encyrtidae). Fla Entomol 85:417–425CrossRefGoogle Scholar
  37. Silva EB, Mouco J, Antunes R, Mendel Z, Franco JC (2009) Mate location and sexual maturity of adult male mealybugs: narrow window of opportunity in a short lifetime. IOBC WRPS Bull 4:3–9Google Scholar
  38. Silva EB, Mendel Z, Franco JC (2010) Can facultative parthenogenesis occur in biparental mealybug species? Phytoparasitica 38:19–21CrossRefGoogle Scholar
  39. Schmidt JM, Smith JJB (1987) Short interval time measurement by a parasitoid wasp. Science 237:903–905PubMedCrossRefGoogle Scholar
  40. Suckling DM (2000) Issues affecting the use of pheromones and other semiochemicals in orchards. Crop Prot 19:677–683CrossRefGoogle Scholar
  41. Tashiro H, Moffitt C (1968) Reproduction in the California red scale. Aonidiella aurantii. II. Mating behaviour and postinsemination female changes. Ann Entomol Soc Am 61:1014–1020Google Scholar
  42. Thornhill R, Alcock J (2001) The evolution of insect mating systems., Inc, San JoseGoogle Scholar
  43. Vet LEM (1983) Host-habitat location through olfactory cues by Leptopilina clavipes (Hartig) (Hym.: Eucoilidae), a parasitoid of fungivorous Drosophila: the influence of conditioning. Neth J Zool 33:225–248CrossRefGoogle Scholar
  44. Watson PJ, Arnqvist G, Stallmann R (1998) Sexual conflict and the energetic costs of mating and mate choice in water striders. Am Nat 151:46–58PubMedCrossRefGoogle Scholar
  45. Waterworth RA, Wright IM, Millar JG (2011) Reproductive biology o the three cosmopolitan mealybug (Hemiptera: Pseudococcidae) species, Pseudococcus longispinus, Pseudococcus viburni, and Planococcus ficus. Ann Entomol Soc Am 104:249–260CrossRefGoogle Scholar
  46. Zada A, Dunkelblum E, Assael F, Harel M, Cojocaru M, Mendel Z (2003) Sex pheromone of the vine mealybug, Planococcus ficus in Israel: Occurrence of a second component in a mass-reared population. J Chem Ecol 29:977–988PubMedCrossRefGoogle Scholar
  47. Zada A, Dunkelblum E, Assael F, Franco JC, Silva EB, Protasov A, Mendel Z (2008) Attraction of Planococcus ficus males to racemic and chiral pheromone baits: flight activity and bait longevity. J Appl Entomol 132:480–489CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2012

Authors and Affiliations

  • Elsa Borges da Silva
    • 1
  • Manuela Branco
    • 1
  • Zvi Mendel
    • 2
  • José Carlos Franco
    • 1
  1. 1.Centro de Estudos Florestais, Instituto Superior de AgronomiaUniversidade Técnica de LisboaLisbonPortugal
  2. 2.Department of EntomologyVolcani Center, AROBet DaganIsrael

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