, Volume 136, Issue 1, pp 29–44 | Cite as

Morphology of the external scent efferent system of Neotropical shield bugs (Hemiptera: Scutelleridae: Pachycorinae)

Original paper


The metathoracic scent glands are characteristic of the true bug suborder Heteroptera and are known to have defensive, sexual and aggregation roles, among others. The morphology of external scent efferent system (ESES) associated with these glands has taxonomic and phylogenetic importance. These structures have been documented previously in four of the eight Scutelleridae subfamilies. In this work we investigate them in detail in 22 genera of the New World subfamily Pachycorinae using scanning electron microscopy. We report five general types of ESES in pachycorine: (1) enlarged evaporatorium and well-developed peritreme; (2) well-developed evaporatorium and poorly developed or obsolete peritreme; (3) reduced evaporatorium and small or obsolete peritreme; (4) reduced evaporatorium on mesepimeron and elongate peritreme; (5) enlarged evaporatorium and elongate peritreme. Our results are in contradistinction to previous accounts of the ESES for the Pachycorinae which have been reported as being poorly developed. The Pachycorinae patterns do not support Schaefer’s hypothesis that there is a correlation between degree of development of the external metathoracic structures and way of life. We also demonstrate that these structures are of systematic importance in the Pachycorinae.


Metathoracic glands Morphology New World Jewel bugs Taxonomy Scanning electron microscopy 



Curators of listed collections are acknowledged for the loan of specimens. We thank to the staff of CMM/UFRGS for the use of facilities and assistance with SEM and to Andressa Paladini, Luiz A. Campos, Petr Kment, Thereza A. Garbelotto and an anonymous reviewer for valuable comments on the manuscript. This study was partially supported by research grants from Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPq (Ed. Universal #470796/2012-0) and by fellowships from Coordenação de Aperfeiçoamento de Pessoal do Ensino Superior – CAPES to L. Weiler (1069496) and from CNPq to K.R. Barão (PDJ #150712/2015-4) and J. Grazia (PQ #305009/2015-0).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

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Supplementary material 1 (DOCX 40 kb)
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Supplementary material 2 (DOCX 30 kb)


  1. Aldrich JR (1988) Chemical ecology of the Heteroptera. Annu Rev Entomol 33:211–238CrossRefGoogle Scholar
  2. Barão KR, Ferrari A, Adami CVK, Grazia J (2016) Morphological diversity of the external thoracic scent efferent system of Carpocorini (Heteroptera: Pentatomidae) with character selection for phylogenetic inference. Zool Anz. doi: 10.1016/j.jcz.2016.08.003
  3. Barcellos A, Eger JE, Grazia J (2014) Scutelleridae. In: Claps LE, Morrone JJ (eds) Biodiversidad de Artrópodos Argentinos, vol 3. Sociedad Entomológica Argentina, Mendoza, pp 409–415Google Scholar
  4. Carapezza A (2009) On some old world Scutelleridae (Heteroptera). Nouv Rev Entomol 25(3):197–212Google Scholar
  5. Carayon J (1955) Quelques caracteres anatomiques des Hemipteres Aradides. Rev Fr Entomol 22:169–180Google Scholar
  6. Carayon J (1971) Notes et documents sur l’appareil odorant métathoracique des Hémiptères. Ann Soc Entomol Fr 7:737–770Google Scholar
  7. Cassis G (1995) A reclassification and phylogeny of the Termatophylini (Heteroptera: Miridae: Deraeocorinae), with a taxonomic revision of the Australian species, and a review of the tribal classification of the Deraeocorinae. Proc Entomol Soc Wash 97:258–330Google Scholar
  8. Cassis G, Schuh RT (2012) Systematics, biodiversity, biogeography, and host associations of the Miridae (Insecta: Hemiptera: Heteroptera: Cimicomorpha). Annu Rev Entomol 57:377–404CrossRefPubMedGoogle Scholar
  9. Cassis G, Vanags L (2006) Jewel bugs of Australia (Insecta, Heteroptera, Scutelleridae). In: Rabitsch W (ed) Hug the bug—for love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss, Denisia 19, pp 275–398Google Scholar
  10. Davidová-Vilímová J, Nejedlá M, Schaefer CW (2000) Dorso-abdominal scent glands and metathoracic evaporatoria in adults of central European Rhopalidae (Hemiptera: Heteroptera), with a discussion of phylogeny and higher systematics. Eur J Entomol 97:213–221CrossRefGoogle Scholar
  11. Drake CJ, Davis NT (1960) The morphology, phylogeny, and higher classification of the family Tingidae, including the description of a new genus and species of the subfamily Vianaidinae (Hemiptera: Heteroptera). Entomol Am 39:1–100Google Scholar
  12. Eger JE (1987) A review of the genus Tiridates Stål (Heteroptera: Pentatomoidea: Scutelleridae). Fla Entomol 70(3):339–350CrossRefGoogle Scholar
  13. Eger JE (1990) Revision of the genus polytes stål (Heteroptera: Scutelleridae). Ann Entomol Soc Am 83(2):115–141CrossRefGoogle Scholar
  14. Eger JE, Barcellos A, Weiler L (2015) Shield bugs (Scutelleridae). In: Panizzi AR, Grazia J (eds) True bugs (Heteroptera) of the neotropics. Springer, Dordrecht, pp 757–788CrossRefGoogle Scholar
  15. Fischer C (2001) Ein Beitragzum Grundmuster, phylogenetischen System und zur Verwandtschaft der Scutelleridae (Heteroptera, Pentatomoidea). Inaugural-Dissertation zur Erlangung der Doktorwürde. Freie Universität, BerlinGoogle Scholar
  16. Gapud V (1991) A generic revision of the subfamily Asopinae with consideration of its phylogenetic position in the family Pentatomidae and superfamily Pentatomoidea (Hemiptera-Heteroptera). Philipp Entomol 8:865–961Google Scholar
  17. Garbelotto TA, Campos LA, Grazia J (2014) Xynocoris, new genus of Ochlerini from central and South America (Hemiptera: Pentatomidae: Discocephalinae). Zootaxa 3869(3):281–305CrossRefGoogle Scholar
  18. Grazia J, Panizzi AR, Greve C, Schwertner CF, Campos LA, Garbelotto TA, Fernandes JAM (2015) Stink bugs (Pentatomidae). In: Panizzi AR, Grazia J (eds) True bugs (Heteroptera) of the neotropics. Springer, Dordrecht, pp 681–756CrossRefGoogle Scholar
  19. Grazia J, Bolze GJ, Barão KR (2016) There and back again: contributions on Pseudevoplitus Ruckes (Heteroptera: Pentatomidae). Zootaxa 4078(1):161–167CrossRefPubMedGoogle Scholar
  20. Hassan SA, Kitching IJ (1993) A cladistic analysis of the tribes of Pentatomidae (Heteroptera). Jpn J Entomol 61:651–669Google Scholar
  21. Kment P (2008) A revision of the endemic Madagascan genus Triplatyx (Hemiptera: Heteroptera: Pentatomidae). Ann Entomol Mus Natl Pragae 48:543–582Google Scholar
  22. Kment P (2015) Two new genera of Madagascan Pentatominae (Hemiptera: Heteroptera: Pentatomidae). Ann Entomol Mus Natl Pragae 55:591–624Google Scholar
  23. Kment P, Vilímová J (2010a) Thoracic scent efferent system of the Tessaratomidae sensu lato (Hemiptera: Heteroptera: Pentatomoidea) with implication to the phylogeny of the family. Zootaxa 2363:1–59Google Scholar
  24. Kment P, Vilímová J (2010b) Thoracic scent efferent system of Pentatomoidea (Hemiptera: Heteroptera): a review of terminology. Zootaxa 2706:1–77Google Scholar
  25. Kment P, Štys P, Vilímová J (2012) Thoracic scent efferent system and exponium of Aphylidae (Hemiptera: Heteroptera: Pentatomoidea), its architecture and function. Eur J Entomol 109:267–279CrossRefGoogle Scholar
  26. Lattin JD (1964) The Scutellerinae of America North of Mexico (Hemiptera: Heteroptera: Pentatomidae). PhD thesis, University of California, BerkeleyGoogle Scholar
  27. Lis JA (2003) Dearcla congoensis, a new species of burrower bugs (Hemiptera: Heteroptera: Cydnidae) from the Democratic Republic of Congo. Polish J Entomol 72:139–143Google Scholar
  28. Matesco VC, Grazia J (2013) Revision of the genus Alkindus Distant (Hemiptera: Heteroptera: Thyreocoridae: Corimelaeninae). Zootaxa 3750(1):057–070CrossRefGoogle Scholar
  29. McDonald FJD, Cassis G (1984) Revision of the Australian Scutelleridae Leach (Hemiptera). Aust J Zool 32:537–572CrossRefGoogle Scholar
  30. Millar JG, McBrien HL, Ho HY, Rice RE, Cullen E, Zalom FG, Üokl A (2002) Pentatomid bug pheromones in IPM: possible applications and limitations. IOBC wprs Bull 25:1–11Google Scholar
  31. Parveen S, Usmani K, Khokhar S, Ramamurthy VV, Vilímová J (2014) External thoracic scent efferent system of Scutelleridae (Hemiptera: Heteroptera). Zool Anz 253:236–258CrossRefGoogle Scholar
  32. Roell T, Campos LA (2015) Candeocoris bistillatus, new genus and new species of Ochlerini from Ecuador (Hemiptera: Heteroptera: Pentatomidae). Zootaxa 4018(4):573–583CrossRefPubMedGoogle Scholar
  33. Schaefer CW (1972) Degree of metathoracic scent-gland development in the Trichophorous Heteroptera (Hemiptera). Ann Entomol Soc Am 65:810–821CrossRefGoogle Scholar
  34. Schaefer CW (2004) Key to genera of new world Alydidae (Hemiptera: Heteroptera). Proc Entomol Soc Wash 106(2):280–287Google Scholar
  35. Schaefer CW, Ahmad I (2008) A revision of Neomegalotomus (Hemiptera: Alydidae). Neotropical Entomol 37(1):030–044CrossRefGoogle Scholar
  36. Schouteden H (1904) Heteroptera. Fam. Pentatomidae. Subfam. Scutellerinae. Genera Insectorum 24:1–98Google Scholar
  37. Schuh RT, Weirauch C, Wheeler WC (2009) Phylogenetic relationships within the Cimicomorpha (Hemiptera: Heteroptera): a total-evidence analysis. Syst Entomol 34:15–48CrossRefGoogle Scholar
  38. Schwertner CF, Grazia J (2015) Less diverse pentatomoid families (Acanthosomatidae, Canopidae, Dinidoridae, Megarididae, Phloeidae, and Tessaratomidae). In: Panizzi AR, Grazia J (eds) True bugs (Heteroptera) of the neotropics. Springer, Dordrecht, pp 821–862CrossRefGoogle Scholar
  39. Staddon BW (1979) The scent glands of Heteroptera. Adv Insect Physiol 14:351–418CrossRefGoogle Scholar
  40. Tsai JF, Rédei D, Yeh GF, Yang MM (2011) Jewel bugs of Taiwan (Heteroptera: Scutelleridae). National Chung Hsing University, TaichungGoogle Scholar
  41. Weirauch C (2006) Metathoracic glands and associated evaporatory structures in Reduvioidea (Heteroptera: Cimicomorpha), with observation on the mode of function of the metacoxal comb. Eur J Entomol 103:97–108CrossRefGoogle Scholar
  42. Weirauch C (2008) Cladistic analysis of Reduviidae (Heteroptera: Cimicomorpha) based on morphological characters. Syst Entomol 33:229–274CrossRefGoogle Scholar
  43. Wood I, McDonald FJD (1984) Revision of the Australian Eysarcoris Group (Hemiptera: Pentatomidae). J Aust Entomol Soc 23:253–264CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  • L. Weiler
    • 1
    • 3
  • K. R. Barão
    • 1
    • 2
  • G. Cassis
    • 3
  • J. Grazia
    • 1
  1. 1.Programa de Pós-Graduação em Biologia Animal, Instituto de BiociênciasUniversidade Federal do Rio Grande do SulPorto AlegreBrazil
  2. 2.Departamento de ZoologiaUniversidade Federal do Rio Grande do SulPorto AlegreBrazil
  3. 3.Evolution and Ecology Research Centre, School of Biological, Earth and Environmental SciencesUniversity of New South WalesSydneyAustralia

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