Eur. J. Entomol. 105 (2): 279-312, 2008 | DOI: 10.14411/eje.2008.038

Morphology of the spermatheca in the Cydnidae (Hemiptera: Heteroptera): Bearing of its diversity on classification and phylogeny

Dominique PLUOT-SIGWALT1, Jerzy A. LIS2
1 Biologie et Évolution des Insectes, École Pratique des Hautes Études et Muséum National d'Histoire Naturelle, 45 rue Buffon, F-75005 Paris, France; e-mail:
2 Department of Biosystematics, Division of Invertebrate Zoology, University of Opole, Oleska 22, PL-45-052 Opole, Poland; e-mail:

Cuticular parts of the spermatheca and associated vaginal structures (chiefly the ring sclerites of the parietovaginal glands) have been examined and compared in 190 cydnid species representing 65 genera and all five subfamilies currently recognized in the family (Amnestinae, Cephalocteinae, Cydninae, Garsauriinae, Sehirinae). Four species belonging to genera formerly included within the Cydnidae (Dismegistus, Parastrachia, Thaumastella, Thyreocoris) were also examined. Morphology of the three main parts of the spermatheca [seminal receptacle (distal bulb), intermediate part (pump apparatus), spermathecal duct] is described. Four main types of spermathecae can be recognized from the distal receptacle and the intermediate part: the amaurocorine type (in Sehirinae: Amaurocorini), amnestine type (in Amnestinae), garsauriine type (in Garsauriinae), and "cydnoid" type (in Cephalocteinae + Cydninae: Cydnini, Geotomini + Sehirinae: Sehirini). No synapomorphy of these types was found which suggests that the currently conceived Cydnidae are not monophyletic. Moreover, out of these four types only the "cydnoid" is typically pentatomoidean due to the presence of an intermediate part usually well delimited by two flanges and having always an unsclerotized flexible zone as well as two internal cuticular structures (septum and fretum) partly obstructing the lumen. The simple tubular amaurocorine type is unusual and aberrant within all Pentatomoidea. The amnestine and garsauriine types display some similarities with taxa outside the Pentatomoidea, especially with some lygaeoid or coreoid spermathecae, mainly in the structure of the intermediate part not delimited proximally (absence of flanges) and devoid of the flexible zone. Within the "cydnoid" type, six spermathecal facies can be characterized principally according to the shape of both the apical reservoir along with the intermediate part, and the differentiations of the spermathecal duct. It has been impossible to find any synapomorphy for all species and for the six facies belonging to the "cydnoid" type of spermatheca. We suggest that the Cydnidae as defined presently are probably a polyphyletic group; moreover its main "cydnoid" branch, called by us Cydnidae sensu stricto (Cephalocteinae + Cydninae + Sehirinae: Sehirini) seems to be relatively recent among the Pentatomoidea. Nishadana and Nishocoris are transferred from Garsauriinae back to Cydninae: Cydnini and the tribe Amaurocorini (Sehirinae) is upgraded to a separate subfamily Amaurocorinae stat. nov. Moreover, we regard the Geotomini and the Sehirini both as non-monophyletic and we indicate that by appending them sensu lato (Geotomini "s. l.", Sehirini "s. l.")

Keywords: Heteroptera, Pentatomomorpha, Pentatomoidea, Cydnidae, Amaurocorinae stat. n., vaginal structures, spermatheca, parietovaginal gland, ring sclerite, fecundation canal, morphology, classification, phylogeny

Received: December 14, 2006; Accepted: December 12, 2007; Revised: December 12, 2007; Published: May 15, 2008Show citation

ACS AIP APA ASA Harvard Chicago IEEE ISO690 MLA NLM Turabian Vancouver
PLUOT-SIGWALT, D., & LIS, J.A. (2008). Morphology of the spermatheca in the Cydnidae (Hemiptera: Heteroptera): Bearing of its diversity on classification and phylogeny. Eur. J. Entomol.105(2), 279-312. doi: 10.14411/eje.2008.038.
Download citation


  1. ADAMS T.S. 2001: Morphology of the internal reproductive system of the male and female of two spotted stink bug, Perillus bioculatus (F.) (Heteroptera: Pentatomidae) and the transfer of products during mating. Invert. Reprod. Devel. 39: 45-53 Go to original source...
  2. AGARWAL S.B. & BAIJAL H.N. 1982: External genitalia of Urostyis pallida Dall. (Pentatomidae: Urostylinae) and phylogenetic significance in family Pentatomidae. J. Entomol. Res. 6: 166-171
  3. AHMAD I. & MOIZUDDIN M. 1980: A new tribe, a new genus and two new species of Cydnidae (Heteroptera: Pentatomoidea) from Pakistan with phylogenetic considerations. Trans. Shikoku Entomol. Soc. 15: 1-9
  4. AHMAD I. & SOUTHWOOD T.R.E. 1964: The morphology of the Alydid abdomen with special reference to the genitalia and its bearing on classification (Heteroptera). Tijdschr. Entomol. 107: 365-378
  5. ANDERSEN N.M. 1982: The Semiaquatic Bugs (Hemiptera, Gerromorpha): Phylogeny, Adaptations, Biogeography, and Classification. Entomonograph 13. Scandinavian Science Press, Klampenborg, 455 pp
  6. ASCHLOCK P.D. 1967: A generic classification of the Orsillinae of the World (Hemiptera - Heteroptera: Lygaeidae). Univ. Calif. Publ. Entomol. 48: 1-82
  7. BERLESE A. 1899: Fenomeni che accompagnano la fecondazione in taluni insetti (Memoria 2). Riv. Patol. Veget. 7[1898]: 1-18
  8. BONHAG P.F. & WICK J.R. 1953: The functional antomy of the male and female reproductive systems of the milkweed bug, Oncopeltus fasciatus (Dallas) (Heteroptera: Lygaeidae). J. Morphol. 93: 177-283 Go to original source...
  9. BRAILOVSKY H. 1984: Revision del genero Latochrimnus Brailovsky (Hemiptera - Heteroptera - Lygaeidae - Lygaeinae). An. Inst. Biol. Univ. Nat. Auton. Mex. (Zool.) 55: 111-128
  10. BRAILOVSKY H. 1987: La tribu Hydarini Stal, en el continente americano con descripcion de dos nuevos generos, una nueva especie y una nueva subspecie (Hemiptera - Heteroptera) (Coreidae). An. Inst. Biol. Univ. Nat. Auton. Mex. (Zool.) 58: 623-650
  11. CARAYON J. 1955: Quelques caracteres anatomiques des Hemipteres Aradides. Rev. Fr. Entomol. 22: 169-180
  12. CARAYON J. 1964: La spermatheque et les voies genitales femelles des Lygaeides Oxycareninae (Heteroptera). Rev. Fr. Entomol. 31: 196-218
  13. COBBEN R.H. 1968: Evolutionary Trends in Heteroptera. Part. I. Eggs, Architecture of the Shell, Gross Embryology and Eclosion. Mededeling n° 151 of the Laboratory of Entomolgy of the agricultural University, Wageningen, 475 pp
  14. COBBEN R.H. 1970: Morphology and taxonomy of intertidal dwarfbugs (Heteroptera: Omaniidae fam. nov.). Tijdschr. Entomol. 113: 61-90
  15. COBBEN R.H. 1980: The Saldidae of the Hawaiian archipelago (Hemiptera: Heteroptera). Pacif. Insects 22: 1-34
  16. COBBEN R.H. 1985: Additions to the Eurasian saldid fauna with a description of fourteen new species (Heteroptera, Saldidae). Tijdschr. Entomol. 128: 215-270
  17. CZAJA J. & LIS J.A. 2002: Four new species of the Afrotropical genus Coleocydnus J.A. Lis (Hemiptera: Heteroptera: Cydnidae). Pol. Pis. Entomol. 71: 273-292
  18. CZAJA J. & LIS J.A. 2003: Coleocydnus inexpectatus, a new Afrotropical species of burrower bugs (Hemiptera: Heteroptera: Cydnidae). Pol. Pis. Entomol. 72: 197-202.
  19. DAVIDOVA-VILIMOVA J. & STYS P. 1980: Taxonomy and phylogeny of West Palearctic Plataspidae (Heteroptera). Stud. CSAV 4: 1-155
  20. DOLLING W.R. 1981: A rationalized classification of the Burrower Bugs (Cydnidae). Syst. Entomol. 6: 61-76 Go to original source...
  21. DUPUIS C. 1955: Les genitalia des Hemipteres Heteropteres (Genitalia externes des deux sexes; Voies ectodermiques femelles). Revue de la morphologie. Lexique de la nomenclature. Index bibliographique analytique. Mem. Mus. Nat. Hist. Nat. (A), Zool. (N.S.) 6: 183-278
  22. DUPUIS C. 1963: Progres recents de l'etude des genitalia des Heteropteres (Etude bibliographique critique). 2e these Doct. Sc., Paris, (Arch. Orig. Centre Doc. CNRS n° 386). Polycopie, 100 pp
  23. DUPUIS C. 1970: 28. Heteroptera. In Tuxen S.L. (ed.): Taxonomist's Glossary of Genitalia in Insect. Munksgaard, Copenhagen, pp. 190-208
  24. DURAI P.S.S. 1987: A revision of the Dinidoridae of the World (Heteroptera: Pentatomoidea). Orient. Insects 21: 163-360 Go to original source...
  25. FISCHER C. 2000: The disc-like organ of the Lestoniidae (Heteroptera: Pentatomoidea), with remarks on lestoniid relationships. Insect Syst. Evol. 31: 201-208 Go to original source...
  26. FREEMAN P. 1947: A revision of the genus Dysdercus Boisduval (Hemiptera, Pyrrhocoridae), excluding the American species. Trans. R. Entomol. Soc. Lond. 98: 373-424 Go to original source...
  27. FROESCHNER R.C. 1960: Cydnidae of the Western Hemisphere. Proc. U. S. Nat. Mus. 111: 337-680 Go to original source...
  28. GAFFOUR-BENSEBBANE C. 1991a: Morphologie des voies genitales ectodermiques des femelles d'Eurygaster austriaca (Schrank, 1776). Bull. Soc. Entomol. Fr. 95[1990]: 209-228
  29. GAFFOUR-BENSEBBANE C. 1991b: Importance des pieces genitales ectodermiques femelles dans la distinction des especes d'Eurygaster (Laporte, 1832). I - Les especes palearctiques (Heteroptera, Scutelleridae). Nouv. Rev. Entomol. (N.S.) 8: 287-305
  30. GAFFOUR-BENSEBBANE C. 1992: Importance des pieces genitales ectodermiques femelles dans la distinction des especes d'Eurygaster (Laporte, 1832). II - Les especes nearctiques (Heteroptera, Scutelleridae). Nouv. Rev. Entomol. (N.S.) 9: 29-46
  31. GAFFOUR-BENSEBBANE C. 1993: Les correlations morphofonctionnelles des organes copulateurs femelles et males chez les Scutelleridae (Heteroptera, Pentatomoidea). Bull. Soc. Entomol. Fr. 98: 313-321
  32. GAFFOUR-BENSEBBANE C. 1994: Les variations morphologiques de l'appareil genital ectodermique des femelles de Scutelleridae (Heteroptera, Pentatomoidea). 1. Les pieces internes. Nouv. Rev. Entomol. (N.S.) 11: 267-281
  33. GAPUD V.P. 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: 865-961
  34. HEMING-VAN BATTUM K. & HEMING B.S. 1986: Structure, function and evolution of the reproductive system in females of Hebrus pusillus and H. rufipes (Hemiptera, Gerromorpha, Hebridae). J. Morphol. 190: 121-167 Go to original source...
  35. HENRY T.J. 1997: Phylogenetic analysis of family groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera). Ann. Entomol. Soc. Am. 90: 275-301 Go to original source...
  36. JACOBS D.H. 1989: A new species of Thaumastella with notes on the morphology, biology and distribution of the two southern African species (Heteroptera: Thaumastellidae). J. Entomol. Soc. Sth Afr. 52: 301-316
  37. KERHZNER I.M. 1976: [New and little known Heteroptera from Mongolia and adjacent regions of the USSR. III. Insects of Mongolia.] 4: 30-86 [in Russian]
  38. KIM H.R. & LEE C.E. 1993: The spermathecae of the Plataspidae from Korea (Heteroptera, Hemiptera). Nature & Life (Korea) 23: 115-120
  39. KIM H.R. & LEE C.E. 1995: Comparative morphology of the spermathecae and the phylogenetic relationships of Korean Cydnidae (Hemiptera, Heteroptera). Russ. Entomol. J. 3[1994]: 41-48
  40. KOCOREK A. & DANIELCZOK-DEMSKA T. 2002: Comparative morphology of the spermatheca within the family Dinidoridae. Eur. J. Entomol. 99: 91-98 Go to original source...
  41. KUMAR R. 1962: Morpho-taxonomical studies on the genitalia and salivary glands of some Pentatomoidea. Entomol. Tidskr. 83: 44-88
  42. KUMAR R. 1965a: Aspect of the morphology of Coreoidea and their value in its higher classification. Proc. R. Soc. Queensl. 76: 27-91
  43. KUMAR R. 1965b: Contribution to the morphology and relationships of Pentatomoidea (Hemiptera: Heteroptera). Part I. Family Scutelleridae. J. Entomol. Soc. Queensl. 4: 41-55 Go to original source...
  44. KUMAR R. 1967: Morphology of the reproductive and alimentary systems of the Aradoidea (Hemiptera), with comments on relationships within the superfamily. Ann. Entomol. Soc. Am. 60: 17-25 Go to original source...
  45. KUMAR R. 1969a: Morphology and relationships of the Pentatomoidea (Heteroptera). III. Natalicolinae and some Tessaratomidae of uncertain position. Ann. Entomol. Soc. Am. 62: 681-695 Go to original source...
  46. KUMAR R. 1969b: Morphology and relationships of the Pentatomoidea (Heteroptera). IV. Oncomerinae (Tessaratomidae). Austr. J. Zool. 17: 553-606 Go to original source...
  47. KUMAR R. 1971: Morphology and relationships of the Pentatomoidea (Heteroptera). 5: Urostylidae. Am. Midl. Nat. 85: 63-73 Go to original source...
  48. KUMAR R & GHAURI M.S.K. 1970: Morphology and relationships of the Pentatomoidea (Heteroptera). 2. World genera of Tessaratomini. Dtsch. Entomol. Z. 17: 1-32 Go to original source...
  49. LEE C.E. & PENDERGRAST J.G. 1983: The spermathecae of New Zealand Aradidae (Hemiptera: Heteroptera). J. Nat. Hist. 17: 113-122 Go to original source...
  50. LEE K.Y., LEE C.E. & PARK H.C. 1989: A comparative study on the spermathecae of the Coreidae from Korea (Heteroptera, Hemiptera). Nature & Life (Korea) 19: 7-14
  51. LESTON D. 1956: The Ethiopian Pentatomoidea (Hemiptera). XXII. On Dismegistus Amyot and Serville (Cydnidae). Proc. R. Entomol. Soc. Lond. (A) 31: 87-94
  52. LINNAVUORI R.E. 1982: Pentatomidae and Acanthosomidae of Nigeria and the Ivory Coast, with remarks on species of the adjacent countries in West and Central Africa. Acta Zool. Fenn. 163: 1-176
  53. LINNAVUORI R.E. 1993: Cydnidae of West, Central and NorthEast Africa (Heteroptera). Acta Zool. Fenn. 192: 1-148
  54. LIS J.A. 1994: A Revision of Oriental Burrower Bugs (Heteroptera: Cydnidae). Upper Silesian Museum, Department of Natural History, Bytom, 349 pp
  55. LIS J.A. 1999: Burrower bugs of the Old World - a catalogue (Hemiptera: Heteroptera: Cydnidae). Genus (Wroclaw) 10: 165-249
  56. LIS J.A. 2002: The mesothoracic wing and its phylogenetic significance in Cydnidae (Hemiptera: Heteroptera). Pol. Pis. Entomol. 71: 43-71
  57. LIS J.A. & HEYNA J. 2001: Metathoracic wings venation in Cydnidae (Hemiptera: Heteroptera) and its bearing on the classification of the family. Ann. Zool. (Warszawa) 51: 429-465
  58. LIS J.A. & HOHOL-KILINKIEWICZ A. 2001: Abdominal trichobothrial pattern and its taxonomic and phylogenetic signifi cance in Cydnidae (Hemiptera: Heteroptera) - II. Amnestinae and Garsauriinae. Ann. Zool. (Warszawa) 51: 467-471
  59. LIS J.A. & HOHOL-KILINKIEWICZ A. 2002: Abdominal trichobothrial pattern and its taxonomic and phylogenetic significance in Cydnidae (Hemiptera: Heteroptera). IV. Cydninae. Pol. Pis. Entomol. 71: 133-143
  60. MAGNIEN P. 1998: Contribution L l'etude du genre Legnotus SchiIdte et description d'une nouvelle espece de Turquie (Heteroptera, Cydnidae). Bull. Soc. Entomol. Fr. 103: 463-473
  61. MAGNIEN P. 2006: A new species from the Near East for the genus Ochetostethus Fieber 1860 (Heteroptera, Cydnidae). In Rabitsch W. (ed.): Hug the Bug - for love of the true bugs. Festschrift zum 70. Geburstag von Ernst Heiss. Denisia 19: 513-516
  62. MALOUF N.S.R. 1933: Studies on the internal anatomy of the stink bug Nezara viridula. Bull. Soc. Entomol. Egypte 17: 96-119
  63. MCDONALD F.J.D. 1966: The genitalia of North American Pentatomoidea (Hemiptera: Heteroptera). Quaest. Entomol. 2: 7-150
  64. MCDONALD F.J.D. 1970: The morphology of Lestonia haustorifera China (Heteroptera: Lestonidae). J. Nat. Hist. 4: 413-417 Go to original source...
  65. MCDONALD F.J.D. 1976: Revision of the genus Trichopepla (Hemiptera: Pentatomidae) in N. America. J. N. Y. Entomol. Soc. 84: 9-22
  66. MCDONALD F.J.D. 1979: A new species of Megaris and the status of the Megarididae McAtee & Malloch and Canopidae Amyot & Serville (Hemiptera: Pentatomoidea). J. N. Y. Entomol. Soc. 87: 42-84
  67. MCDONALD F.J.D & CASSIS G. 1984: Revision of the Australian Scutelleridae Leach (Hemiptera). Austr. J. Zool. 32: 537-572 Go to original source...
  68. MOIZUDDIN M. & AHMAD I. 1990: A revision of Cydnidae (Hemiptera: Pentatomomorpha: Pentatomoidea) from Indo-Pakistan area with a cladistic analysis of the included genera. Orient. Insects 24: 305-354 Go to original source...
  69. MONTEITH G.B. 1997: Revision of the Australian flat bugs of the subfamily Mezirinae (Insecta: Hemiptera: Aradidae). Mem. Queensl. Mus. 41: 1-169
  70. MOULET P. 1993: Structures meconnues dans la spermatheque d'Heteropteres Coreoidea palearctiques. Ann. Soc. Entomol. Fr. (N.S.) 29: 159-172
  71. O'DONNELL J.E. 2000: Notes on the phylogenetic relationships of Neotropical Antillocorini, with description of three new species (Heteroptera: Rhyparochromidae: Rhyparochrominae). J. N. Y. Entomol. Soc . 108: 114-129 Go to original source...
  72. PENDERGRAST J.G. 1957: Studies on the reproductive organs of the Heteroptera with consideration of their bearing on classification. Trans. R. Entomol. Soc. Lond. 109: 1-63 Go to original source...
  73. PERICART J. 1990: Hemipteres Saldidae et Leptopodidae d'Europe occidentale et du Maghreb. Faune de France 77. Federation francaise des Societes de Sciences naturelles, Paris, 238 pp
  74. PLUOT D. 1970: La spermatheque et les voies genitales femelles des Pyrrhocorides (Hemiptera). Ann. Soc. Entomol. Fr. (N.S.) 6: 777-807
  75. POLIVANOVA E.N. 1959: [Importance des caracteres anatomiques pour la systematique et la determination des correlations phylogenetiques chez les Heteropteres, Pentatomoidea]. Trudy Inst. Morph. Zhivotn. Severtsova 27: 201-220 [in Russian]
  76. PUTCHKOVA L.V. 1959: [Traits generaux de la structure des oeufs et de l'appareil genital femelle des Heteropteres L scutellum]. Praci Inst. Zool. Acad. Ukrain. 15: 117-125 [in Ukrainian, Russian abstr.]
  77. RAMAMURTY P.S. 1969: Histological studies of the internal organs of reproduction in Nezara viridula Fabr. (Pentatomidae - Heteroptera, Insecta). Zool. Anz. 183: 119-139
  78. RAMAMURTY P.S. & MEDHI W.P. 1970: The internal reproductive organs of Cydnus indicus Westw. A histological study (Cydnidae, Heteroptera, Insecta). La Cellule 68: 97-106
  79. RIEGER C. 1997: Ergaenzungen zur Faunistik und Systematik einiger Wanzen in Baden-Wuerttemberg (Insecta, Heteroptera) II. Carolinea 55: 43-48
  80. RIZZOTTI VLACH M. 2000: Una nuova specie di Ochetostethus della Sardegna e note sulla distribuzione del genere in Italia (Heteroptera, Cydnidae). Nouv. Rev. Entomol. (N.S.) 17: 69-76
  81. ROLSTON L.H. & MCDONALD F.J.D. 1979: Keys and diagnoses for the families of Western Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae (Hemiptera). J. N. Y. Entomol. Soc . 87: 189-207
  82. SCHAEFER C.W. 1981: The sound-producing structures of some primitive Pentatomoidea (Hemiptera: Heteroptera). J. N. Y. Entomol. Soc. 88: 230-235
  83. SCHAEFER C.W. 1988: The food plands of some "primitive" Pentatomoidea (Hemiptera: Heteroptera). Phytophaga 2: 19-45
  84. SCHAEFER C.W. 1993a: The Pentatomomorpha (Hemiptera: Heteroptera): an annotated outline of its systematic history. Eur. J. Entomol. 90: 105-122
  85. SCHAEFER C.W. 1993b: Notes on the morphology and family relationships of Lestoniidae (Hemiptera: Heteroptera). Proc. Entomol. Soc. Wash. 95: 453-456
  86. SCHAEFER C.W., DOLLING W.R. & TACHIKAWA S. 1988: The shieldbug genus Parastrachia and its position within the Pentatomoidea (Insecta: Hemiptera). Zool. J. Linn. Soc. 93: 283-311 Go to original source...
  87. SCHMITZ G. 1976: La faune terrestre de l'ile de Sainte-Helene. 3. Fam. Cydnidae. Ann. Mus. R. Afr. Centr. (Zool.) n° 215: 353-367
  88. SCHUH R.T. & SLATER J.A. 1995: True Bugs of the World (Hemiptera: Heteroptera), Classification and Natural History. Cornell University Press, Ithaca and London, 336 pp
  89. SCUDDER G.G.E. 1959: The female genitalia of the Heteroptera: morphology and bearing on classification. Trans. R. Entomol. Soc. Lond. 111: 405-467 Go to original source...
  90. SCUDDER G.G.E. 1962: New Heterogastrinae (Hemiptera) with a key to the genera of the World. Opusc. Entomol. 27: 117-127
  91. SCUDDER G.G.E. 1965: Neacoryphus Scudder, a new genus of Lygaeinae (Hemiptera, Lygaeidae). Proc. Entomol. Soc. Br. Columb. 62: 34-37
  92. SERVADEI A. 1964: Il valore tassonomico delle spermateche degli Emitteri Eterotteri (fam. Pentatomidae & Acanthosomatidae). Atti Acad. Naz. Ital. Entomol. 11: 58-86
  93. SLATER J.A. 1950: An investigation of the female genitalia as taxonomic characters in the Miridae (Hemiptera). Iowa State Coll. J. Sci. 25: 1-81
  94. SLATER J.A. & ASHLOCK P.D. 1980: On the genus Camptocoris Puton with descriptions of two new species from South Africa (Hemiptera: Lygaeidae). J. Entomol. Soc. Sth Afr. 43: 53-63
  95. SLATER J.A. & CARAYON J. 1963: Ethiopian Lygaeidae IV: a new predatory Lygaeid from Africa with a discussion of its biology and morphology (Hemiptera: Heteroptera). Proc. R. Entomol. Soc. Lond. (A) 38: 1-11
  96. SLATER J.A. & SWEET M.E. 1977: The genus Plinthisus Stephens in the australian Region (Hemiptera, Lygaeidae). Entomol. Scand. 8: 109-154 Go to original source...
  97. STEHLIK J.L. 1965a: Mission zoologique de l'I.R.S.A.C. en Afrique orientale (P. Basilewsky - N. Leleup, 1957). Pyrrhocoridae (Het.). Acta Mus. Morav. 50: 211-252.
  98. STEHLIK J.L. 1965b: Pyrrhocoridae and Largidae collected by E.S. Brown on Solomon Islands (Heteroptera). Acta Mus. Morav. 50: 253-292
  99. STYS P. 1962: Morphology of the abdomen and female ectodermal genitalia of the trichophorous Heteroptera and bearing on their classification. Trans. 11th Int. Congr. Entomol Vienna 1[1961]: 37-43
  100. STYS P. 1964: Thaumastellidae - a new family of Pentatomoid Heteroptera. Acta Soc. Entomol. Cechoslov. 61: 238-253
  101. STYS P. 1966: Morphology of the wings, abdomen and genitalia of Phaenacantha australiae Kirk. (Heteroptera, Colobatristidae) and notes on the phylogeny of the family. Acta Entomol. Bohemoslov. 63: 266-280
  102. STYS P. & DAVIDOVA J. 1979: Taxonomy of Thyreocoris (Heteroptera, Thyreocoridae). Annot. Zool. Bot. 134: 1-40
  103. SWEET M.H. & SCHAEFER C.W. 2002: Parastrachiinae (Hemiptera: Cydnidae) raised to family level. Ann. Entomol. Soc. Am. 95: 441-448 Go to original source...
  104. VAVRINOVA I. 1988: Spermathecae of Central European species of the families Rhopalidae, Alydidae and Coreidae (Heteroptera: Coreoidea). Cas. Morav. Mus. Brne 73: 203-215
  105. WAGNER E. 1963: Untersuchungen ueber den taxonomischen Wert des Baues der Genitalien bei den Cydnidae (Hem. Het.). Acta Entomol. Mus. Nat. Pragae 35: 73-115

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY), which permits use, distribution, and reproduction in any medium, provided the original publication is properly cited. No use, distribution or reproduction is permitted which does not comply with these terms.