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. 2008 Jan 2;13(4):465–501. doi: 10.1111/j.1365-3113.1988.tb00256.x

Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea)

JAMES B WOOLLEY 1,✉
PMCID: PMC7169575  PMID: 32327872

Abstract

Abstract. A data set consisting of twenty‐eight anatomical characters scored for twenty‐eight terminal taxa representing the world fauna of Signiphoridae was analysed using parsimony and compatibility methods. The Coccophaginae (Aphelinidae) and the Azotinae (Aphelinidae) were used as outgroups to establish polarity of character state changes. Relationships of Signiphoridae to other Chalcidoidea are discussed. Several multistate characters were treated in the parsimony analyses either as unordered or as ordered into transformation series using additive binary coding, which in some cases drastically reduced the number of equally parsimonious solutions. Monophyly of Signiphoridae is supported by seven synapomorphies. Four genera, Chartocerus, Thysanus, Clytina and Signiphora, are recognized within Signiphoridae based on synapomorphies. Rozanoviella syn.n. and Kerrichiella syn.n. are synonymized under Signiphora. Species of Signiphora are further assigned to four species groups, three of which are demonstrably monophyletic. Nine species or subspecies are transferred to Chartocerus from Signiphora (australicus comb.n., australiensis comb.n., australiensis orbiculatus comb.n., beethoveni comb.n., corvinus comb.n., funeralis comb.n., reticulata comb.n., ruskini comb.n., thusanoides comb.n.), one species to Thysanus from Signiphora (melancholicus comb.n.), and one species to Signiphora from Kerrichiella (coleoptratus comb.n.). A key to genera of Signiphoridae and species groups of Signiphora is presented. A diagnosis, relevant nomenclatural history, and a list of included species are given for each genus and species group, and the biology and distribution of each is summarized.

References

  1. Agarwal, M.M. (1963) Matritia kerrichi n.sp. (Hymenoptera: Thysanidae) an endoparasite of Eriococcus greeni Newst. recorded from Aligarh (India). Zeitschrift fiir Parasitenkunde, 22, 389–393. [Google Scholar]
  2. Agekyan, N.G. (1968) Signiphora merceti Malen. (Hymenoptera, Chalcidoidea) ‐ a parasite of Hemiberlesia rapax (Comst.) in Adzharia. Entomological Review, 47, 484–486. [Google Scholar]
  3. Ashmead, W.H. (1880) Orange Insects. A Treatise on the Injurious and Beneficial Insects Found on the Orange Trees of Florida. Ashmead Bros., Jacksonville , Florida . [Google Scholar]
  4. Ashmead, W.H. (1900) Genera of the Encyrtinae. Subfamily III ‐ Signiphorinae. Proceedings of the U.S. National Museum, 22, 409–412. [Google Scholar]
  5. Ashmead, W.H. (1904) Classification of the chalcid flies or the superfamily Chalcidoidea with descriptions of new species in the Carnegie Museum, collected in South America by Herbert J. Smith. Memoirs of the Carnegie Museum, 1, xi+225–551. [Google Scholar]
  6. Blanchard, E.E. (1936) Apuntes sobre Calcidoideos Argentinos, Nuevos y Conocidos. Revista Sociedad Entomologica Argentina, 8, 1–32. [Google Scholar]
  7. Brèthes, J. (1913) Himenópteros de la America Meridional. Anales de Museo Nacional de Historia Natural de Buenos Aires, 24, 35–165. [Google Scholar]
  8. Brèthes, J. (1914) Les ennemies de la ‘Diaspis pen‐tagona’ dans la Republique Argentine. Nunquam Otiosus , pp. 1–16.
  9. Clausen, C.P. (1924) The parasites of Pseudococcus maritimus (Ehrhorn) in California (Hymenoptera, Chalcidoidea). Part II. Biological studies and life histories. University of California Publications, Technical Bulletin, Entomology, 3, 253–292. [Google Scholar]
  10. Compere, H. (1955) A systematic study of the genus Aphytis Howard (Hymenoptera, Aphelinidae) with descriptions of new species. University of California, Publications in Entomology, 10, 271–320. [Google Scholar]
  11. Compere, H. & Annecke, D.P. (1961) Descriptions of parasitic Hymenoptera and comments (Hymenopt.: Aphelinidae, Encyrtidae, Eulophidae). Journal of the Entomological Society of Southern Africa, 24, 17–71. [Google Scholar]
  12. Crawford, J.C. (1913) Descriptions of new Hymenoptera, No. 8. Proceedings of the U.S. National Museum, 46, 343–352. [Google Scholar]
  13. de Dalla Torre, C.G. (1898) Catalogus Hymenop‐terorum hucusque descriptorum systemicus et syn‐onymicus. Volumen V: Chalcididae et Proctotrupidae. Sumptibus Guilelmi Engelmann, Lipsiae . [Google Scholar]
  14. Darling, CD. & Johnson, N.F. (1984) Synopsis of Nearctic Azotinae (Hymenoptera: Aphelinidae). Proceedings of the Entomological Society of Washington, 86, 555–562. [Google Scholar]
  15. DeBach, P. (1953) Thysanus flavopalliatus (Ashm.) parasitic on Compendia bifasciata How. on California red scale. Journal of Economic Entomology, 36, 1112. [Google Scholar]
  16. DeBach, P. , Kennett, C.E. & Pence, R.J. (1958) Species of Thysanus as primary parasites. Journal of Economic Entomology, 51, 114–115. [Google Scholar]
  17. De Santis, L. (1968) Nomenclatura y clasificación de la familia Signiphoridae (Hymenoptera: Chalcidoidea). Revista de la Facultad de Agronomia, 44, 7–16. [Google Scholar]
  18. De Santis, L. (1973) Nota sobre signifóridos de la Republica Argentina (Hymenoptera: Chalcidoidea). Revista de la Facultad de Agronomia, 49, 143–153. [Google Scholar]
  19. De Santis, L. (1979) Catálogo de los Himenópteros Calcidoideos de América al Sur de los Estados Unidos. Publicación Especial, Prov. de Buenos Aires, Comisión de Investigaciones Cientificas, La Plata . [Google Scholar]
  20. De Santis, L. (1981) Catálogo de los Himenópteros Calcidoídeos de América al Sur de los Estados Unidos ‐ Primer Suplemento. Revista Peruana de Entomologia, 24, 1–38. [Google Scholar]
  21. Domenichini, G. (1953) Studio sulla morfologia dell'addome degli Hymenoptera Chalcidoidea. Bollettino di Zoologia Agraria e di Bachicoltura, 19, 1–117. [Google Scholar]
  22. Domenichini, G. (1954) Sulla morfologia e posizione sistematica dei Thysanidae (=Signiphoridae) (Hym. Chalcidoidea). Bollettino di Zoologia Agraria e di Bachicoltura, 20, 93–110. [Google Scholar]
  23. Domenichini, G. (1955) Variabilita dei caratteri e nuova diagnosi di un Tisanidae (Hym. Chalcidoidea) con la descrizione di una nuova specie. Annali della Facolta di Agraria (Nuova Serie), 4, 25–42. [Google Scholar]
  24. Donoghuc, M.J. & Cantino, P.D. (1984) The logic and limitations of the outgroup substitution approach to cladistic analysis. Systematic Botany, 9, 192–202. [Google Scholar]
  25. Dozier, H.L. (1933) Miscellaneous notes and descriptions of chalcidoid parasites (Hymenoptera). Proceedings of the Entomological Society of Washington, 35, 85–100. [Google Scholar]
  26. Erdös, J. (1957) Recentiores observationes entomo‐coenologicae in Phragmite communi Trin. Allat-tani Kozlemenyek, 46, 49–65. [Google Scholar]
  27. Estabrook, G.F. (1979) Some concepts for the estimation of evolutionary relationships in systematic botany. Systematic Botany, 3, 146–158. [Google Scholar]
  28. Estabrook, G.F. , Johnson, C.S., Jr & McMorris, F.R. (1976a) A mathematical foundation for the analysis of character compatibility. Mathematical Biosciences, 23, 181–187. [Google Scholar]
  29. Estabrook, G.F. , Johnson, C.S., Jr & McMorris, F.R. (1976b) An algebraic analysis of cladistic characters. Discrete Mathematics, 16, 141–147. [Google Scholar]
  30. Farris, J.S. (1970) Methods for computing Wagner trees. Systematic Zoology, 19, 83–92. [Google Scholar]
  31. Farris, J.S. (1982) Outgroups and parsimony. Systematic Zoology, 31, 328–334. [Google Scholar]
  32. Felsenstein, J. (1984) The statistical approach to inferring evolutionary trees and what it tells us about parsimony and compatibility Cladistics: Perspectives on the Reconstruction of Evolutionary History (ed. By Duncan T. and Stuessy T.F.), pp. 169–171. Columbia University Press, New York . [Google Scholar]
  33. Ferrière, C. (1953) Encyrtides paléartiques (Hym. Chalcidoidea) Nouvelle table des generes avec notes et synonymies. Mitteilungen Schweizerischen Entomologischen Gessellschaft, 26, 1–45. [Google Scholar]
  34. Foerster, A. (1856) Hymenopterologische Studien. Vol. II, Chalcidiae und Proctotrupii. Verlag von Ernst ter Meer, Aachen . [Google Scholar]
  35. Foester, A. (1878) Kleine Monographien Para‐sitischer Hymenopteren. Verhandlungen Natur-historischen der Vereins der Preussischen Rheinlande und Westfalens, 35, 42–82. [Google Scholar]
  36. Gauld, I.D. & Mound, L.A. (1982) Homoplasy and delineation of holophyletic genera in some insect groups. Systematic Entomology, 7, 73–86. [Google Scholar]
  37. Gibson, G.A.P. (1985) Some pro‐ and mesothoracic characters important for phylogenetic analysis of Hymenoptera, with a review of terms used for structures. Canadian Entomologist, 117, 1395–1443. [Google Scholar]
  38. Gibson, G.A.P. (1986) Mesothoracic skeletomuscula‐ture and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). Canadian Entomologist, 118, 691–728. [Google Scholar]
  39. Girault, A. A. (1913a) A systematic monograph of the chalcidoid Hymenoptera of the subfamily Sig‐niphorinae. Proceedings of the U.S. National Museum, 45, 189–233. [Google Scholar]
  40. Girault, A.A. (1913b) Australian Hymenoptera Chalcidoidea. IV. The Family Eulophidae with descriptions of new genera and species. Memoirs of the Queensland Museum, 2, 140–296. [Google Scholar]
  41. Girault, A.A. (1913c) A new Signiphora from Queensland, Australia (Hym.). Entomological News, 24, 166–167. [Google Scholar]
  42. Girault, A.A. (1915) Australian Hymenoptera Chalcidoidea. VII. The Family Encyrtidae with descriptions of new genera and species. Memoirs of the Queensland Museum, 4, 1–184. [Google Scholar]
  43. Girault, A.A. (1916a) New Encyrtidae from North America. Psyche, 23, 41–42. [Google Scholar]
  44. Girault, A.A. (1916b) Descriptiones Hymenop‐terorum Chalcidoidicorum Variorum cum Obser‐vationibus II. Entomological News, 27, 401. [Google Scholar]
  45. Girault, A. A. (1917) Some new Mymaridae and a new Signiphora in the collection of the Zoological Museum, Berlin. Societas Entomologica, 32, 19–20. [Google Scholar]
  46. Girault, A. A. (1921) Miscellaneous species of chalcid‐flies from Australia (Hymenoptera, Chalcididae). Insecutor Inscitiae Menstruus, 9, 186–189. [Google Scholar]
  47. Girault, A.A. (1929) Notes on, and descriptions of chalcid wasps in the South Australian Museum. Concluding Paper. Transactions Royal Society South Australia, 53, 309–346. [Google Scholar]
  48. Girault, A.A. (1932) New Lower Hymenoptera from Australia and India. Brisbane, Australia. 6 pages. [Privately printed, reprinted in Gordh, G., Menke, A.S., Dahms, E.C. and Hall, J.C. (1979) The Privately Printed Papers of A.A. Girault, Memoirs of the American Entomological Institute, Number28, 400 pages].
  49. Girault, A.A. (1933) Some Beauties Inhabitant not of the Boudoirs of Commerce but of Nature's Bosom ‐ New Insects. Brisbane, Australia. 2 pages, [privately printed, reprinted in Gordh et al., 1979, op.cit.].
  50. Girault, A.A. (1935) Microhymenoptera Australien‐sis Nova, mostly Chalcididae. Sidney, Australia. 4 pages, [privately printed, reprinted in Gordh et al., 1979, op.cit.] .
  51. Gordh, G. (1979) Family Encyrtidae Catalog of Hymenoptera in America North of Mexico, Vol. I, Symphyta and Apocrita (Parasitica) (ed. By Krombein K. V., Hurd P. D., Jr, Smith D. R. and Burks B. D.), pp. 890–967. Smithsonian Institution Press, Washington , D.C . [Google Scholar]
  52. Hayat, M. (1970) Studies on the genera of the family Signiphoridae (Hymenoptera: Chalcidoidea) recorded from India. Entomophaga, 15, 387–399. [Google Scholar]
  53. Hayat, M. (1976) Some Indian species of Chartocerus (Hym.: Chalcidoidea: Signiphoridae). Oriental Insects, 10, 161–164. [Google Scholar]
  54. Hayat, M. (1983) The genera of Aphelinidae (Hymenoptera) of the world. Systematic Entomology, 8, 63–102. [Google Scholar]
  55. Hayat, M. & Verma, M. (1980) A catalogue of the Indian Signiphoridae (Insecta: Hymenoptera: Chalcidoidea). Journal of the Bombay Natural History Society, 76, 481–485. [Google Scholar]
  56. Hecht, M.K. & Edwards, J.S. (1977) The methodology of phylogenetic inference above the species level Major Patterns in Vertebrate Evolution (ed. By Hecht M. K., Goody P. C. and Hecht B. M.), pp. 3–51. NATO Advanced Study Institute Series A, Volume 14 Plenum Press, New York . [Google Scholar]
  57. Hennig, W. (1966) Phylogenetic Systematics. University of Illinois Press, Urbana . [Google Scholar]
  58. Heppner, J.B. & Lamas, G. (1982) Acronyms for world museum collections of insects, with an emphasis on Neotropical Lepidoptera. Bulletin of the Entomological Society of America, 28, 305–315. [Google Scholar]
  59. Heraty, J.M. & Darling, D.C. (1984) Comparative morphology of the planidial larvae of Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea). Systematic Entomology, 9, 309–328. [Google Scholar]
  60. Howard, L.O. (1894) The hymenopterous parasites of the California red scale. Insect Life (USDA), 4, 227–236. [Google Scholar]
  61. Howard, L.O. (1895) Revision of the Aphelininae of North America. Technical Series No. 1, 55 pp., U.S. Department of Agriculture, Division of Entomology, USDA.
  62. International Commission on Zoological Nomenclature (1979) Opinion 1143. Kerrichiella Rosanov, 1965 (Hymenoptera): designation of a type species under the plenary powers. Bulletin of Zoological Nomenclature, 36, 130–131. [Google Scholar]
  63. Kerrich, G.J. (1953) Report on Encyrtidae associated with mealybugs on cacao in Trinidad and on some other species related thereto. Bulletin of Entomological Research, 44, 789–810. [Google Scholar]
  64. Khasef, M.A. (1953) Sur la presence de formations particulieres dans les mandibules de Lariophagus distinguendus Foerst. (Hym. Pteromalidae). Bulletin de la Sociiti Entomologique de France, 58, 141–143. [Google Scholar]
  65. Kluge, A.G. & Farris, J.S. (1969) Quantitative phylogenetics and the evolution of Anurans. Systematic Zoology, 18, 1–32. [Google Scholar]
  66. Kurdjumov, N.V. (1916) New genus and species of Aphelininae (Chalcidoidea). Zhurnal Prikladnoi Entomologii (Kiev), 1, 80–81. [Google Scholar]
  67. LaSalle, J. (1987) New World Tanaostigmatidae (Hymenoptera, Chalcidoidea). Contributions of the American Entomological Institute, 23, 1–181. [Google Scholar]
  68. LaSalle, J. & Noyes, J.S. (1985) New family placement for the genus Cynipencyrtus (Hymenoptera: Chalcidoidea: Tanaostigmatidae). Journal of the New York Entomological Society, 93, 1261–1264. [Google Scholar]
  69. Le Quesne, W.J. (1972) Further studies based on the uniquely derived character concept. Systematic Zoology, 21, 281–288. [Google Scholar]
  70. Lundberg, J.G. (1972) Wagner networks and ancestors. Systematic Zoology, 18, 1–32. [Google Scholar]
  71. Maddison, W.P. , Donoghue, M.J. & Maddison, D.R. (1984) Outgroup analysis and parsimony. Systematic Zoology, 33, 83–103. [Google Scholar]
  72. Malenotti, E. (1916) ‘Signiphora merceti’ Malen. n.sp. Redia, 12, 181–182. [Google Scholar]
  73. Malenotti, E. (1918) I nemici naturali della ‘Bianci‐Rossa’(Chrysomphalus dictyospermi Morg.). Redia, 13, 17–53. [Google Scholar]
  74. Matta, V. (1979) Enemigos naturales de las con‐chuelas blancas del olivo en el Valle de Azapa, Arica‐Chile. Idesia, 5, 231–242. [Google Scholar]
  75. Mercet, R.G. (1912) Los enemigos de los parasitos de las plantas. Los Afelininos. Trabajos del Museo de Ciencias Naturales Madrid, 10, 1–306. [Google Scholar]
  76. Mercet, R.G. (1916) Signiphorinos de Espana (Himenopteros Chalcídidos). Boletin de la Real Sociedad Espanola de Historia Natural, 16, 519–533. [Google Scholar]
  77. Mercet, R.G. (1917) Revisión de los Signiphorinos de España. Revista de la Real Academia de Ciencias Exactas, Fisicas y Naturales, de Madrid, 16, 1–11. [Google Scholar]
  78. Mercet, R.G. (1929) Los Afelínidos de España. Revista de Biologia Forestaly Limnologia. Año I, Serie B, Núm. 1, 1–28. [Google Scholar]
  79. de Motschulsky, N. (1859) IV. Entomologie appli‐quee. Insectes utiles et nuisibles. Etudes Entomologiques, 8, 169–174. [Google Scholar]
  80. Nikol'skaya, M.N. (1950) Representatives of the family Signiphoridae (Hymenoptera: Chalcidoidea) in the fauna of the U.S.S.R. Doklady Akademii Nauk SSSR, 75, 319–321. [In Russian]. [Google Scholar]
  81. Nikol'skaya, M.N. & Yashnosh, V.A. (1966) Aphelinidae of the European Part of the U.S.S.R. and the Caucasus (Chalcidoidea, Aphelinidae). Opredeliteľ Faune SSR, No. 91 , 296 pp. [In Russian].
  82. Peck, O. (1951) Family Thysanidae Hymenoptera of America North of Mexico ‐ Synoptic Catalog (ed. By Muesebeck C. F. W., Krombein K. V. and Townes H. K.), pp. 472–474 U.S. Department of Agriculture, Agricultural Monograph, 2. [Google Scholar]
  83. Quezada, J.R. , DeBach, P. & Rosen. D. (1973) Biological and taxonomic studies of Signiphora borin‐quensis, new species (Hymenoptera: Signiphoridae), a primary parasite of diaspine scales. Hilgardia, 41, 543–604. [Google Scholar]
  84. Richards, O.W. (1935) Two new parasites of aculeate Hymenoptera from Trinidad. Stylops, 4, 131–133. [Google Scholar]
  85. Riek, E.F. (1970) Hymenoptera. Insects of Australia (ed. By CSIRO, Division of Entomology), pp. 867–959. Melbourne University Press. [Google Scholar]
  86. Rohlf, F.J. (1982) Consensus indices for comparing classifications. Mathematical Biosciences, 59, 131–144. [Google Scholar]
  87. Rozanov, L. V. (1965) Review of the genera of parasitic Hymenoptera of the family Signiphoridae (Hymenoptera, Chalcidoidea). Entomological Review, 44, 508–515. [Google Scholar]
  88. Rust, E.W. (1913) New Peruvian parasites from Hemichionaspis minor (Hym.). Entomological News, 24, 160–165. [Google Scholar]
  89. Schauff, M.E. (1984) The Holarctic genera of Mymaridae (Hymenoptera: Chalcidoidea). Memoirs of the Entomological Society of Washington, 12, 1–67. [Google Scholar]
  90. Schmiedeknecht, O. (1909) Hymenoptera. Family Chalcididae Genera Insectorum (ed. By Wyts‐man P.), Fasc. 97, 550 pp. Verteneuil and Desmet, Bruxelles . [Google Scholar]
  91. Snodgrass, R.E. (1941) The male genitalia of the Hymenoptera. Smithsonian Miscellaneous Collections, 99, 1–86. [Google Scholar]
  92. Sokal, R.R. & Sneath, P.H.A. (1963) Principles of Numerical Taxonomy. W. H. Freeman, San Francisco . [Google Scholar]
  93. Strauch, J.G. (1984) Use of homoplastic characters in compatibility analysis. Systematic Zoology, 33, 167–177. [Google Scholar]
  94. Rao Subba , B.R. (1957) Some new species of Indian Hymenoptera. Proceedings of the Indian Academy of Sciences Section B, 46, 376–390. [Google Scholar]
  95. Rao Subba , B.R. (1974) The genera of Signiphoridae (Hymenoptera) with description of a new genus. Bulletin of Entomological Research, 64, 525–531. [Google Scholar]
  96. Sugonyaev, E. (1968) New chalcid species (Hymenoptera, Chalcidoidea) parasitic on jumping plant lice and scale insects (Homoptera, Psyllodea, Coc‐cidae) from the deserts and wastelands of Soviet Central Asia and Kazakhstan. Entomological Review, 47, 361–370. [Google Scholar]
  97. Swofford, D.L. (1985) PAUP‐Phylogenetic Analysis using Parsimony, Version 2.4 Unpublished user's manual, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820.
  98. Throckmorton, L.H. (1965) Similarity versus relationship in Drosophila . Systematic Zoology, 14, 221–236. [PubMed] [Google Scholar]
  99. Timberlake, P.H. (1924) The parasites of Pseudococ‐cus maritimus (Ehrhorn) in California (Hymenoptera, Chalcidoidea). Part I. Taxonomic studies. University of California Publications, Technical Bulletins, Entomology, 3, 223–251. [Google Scholar]
  100. Trjapitsyn, V.A. (1968) Structural features of the female abdomen of the Encyrtidae (Hymenoptera) and their taxonomic significance. Entomological Review, 47, 177–284. [Google Scholar]
  101. Trjapitsyn, V.A. (1977) The characteristics of the morphology of adult encyrtids (Hymenoptera, Chalcidoidea, Encyrtidae) and their systematic significance. Trudy Vsesoyuznogo Entomo-logischeskogo Obshchestva, 58, 145–199. [In Russian]. [Google Scholar]
  102. Walker, F. (1840) Descriptions of British Chalcidites. Annals and Magazine of Natural History, 4, 232–236. [Google Scholar]
  103. Walker, F. (1872) Notes on Chalcidiae. Part VII. E. W. Janson, London . [Google Scholar]
  104. Woolley, J.B. (1986) Signiphoridae or Thysanidae? A review of a problem in family‐level nomenclature (Hymenoptera: Chalcidoidea). Bulletin of the Entomological Society of America, 32, 91–96. [Google Scholar]
  105. Woolley, J.B. & Vet, L.E.M. (1981) Postovipositional web‐spinning behavior in a hyperparasite, Sig‐niphora coquilletti Ashmead (Hymenoptera: Signiphoridae). Netherlands Journal of Zoology, 31, 627–633. [Google Scholar]
  106. Yashnosh, V.A. (1976) Classification of the parasitic Hymenoptera of the family Aphelinidae (Chalcidoidea). Entomological Review, 55, 114–120. [Google Scholar]
  107. Yoshimoto, CM. (1965) The Hawaiian Thysaninae. Pacific Insects, 7, 703–704. [Google Scholar]

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