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2012 - Professor Gerry Cassis, Anna Namyatova, Nikolai Tatarnic and Celia Symonds, University of New South Wales, Sydney
15 February 2002
The Pentatomoidea is the most diverse superfamily of pentatomomorphan bugs and includes the stinkbugs, shieldbugs, tortoise bugs, burrower bugs and jewel bugs. The superfamily comprises 14 families: Acanthosomatidae, Aphylidae, Canopidae, Cydnidae, Dinidoridae, Lestoniidae, Megarididae, Pentatomidae, Phloeidae, Plataspidae, Scutelleridae, Tessaratomidae, Thaumastellidae and Urostylidae.
The pentatomoid families present in Australia are: Acanthosomatidae, Aphylidae, Cydnidae, Dinidoridae, Lestoniidae, Pentatomidae, Plataspidae, Scutelleridae and Tessaratomidae. Monteith pers. comm. has indicated that the Urostylidae do not occur in Australia, that previous records are in error (Woodward et al. 1970; Carver et al. 1991). Thus, in this Catalogue they are excluded from the Australian heteropteran fauna. The Canopidae, Megarididae, Phloeidae, Thaumastellidae and Urostylidae are not represented in Australia.
The Pentatomoidea have been widely accepted as a higher taxon within the Pentatomomorpha (Pendergrast 1957; Leston 1958; Scudder 1959; China & Miller 1959; McDonald 1966; Gapud 1991; Carver et al. 1991; Schuh & Slater 1995). Much of the literature is summarised by Göllner-Scheiding (1990), Stonedahl & Dolling (1991), Schaefer (1993) and Henry (1997).
The Thaumastellidae are restricted to Afrotropical and Palaearctic Regions and are the only taxon whose position in the Pentatomoidea is debatable. This family was originally placed in the Lygaeidae and subsequently transferred to the Pentatomoidea (at family rank) by Štys (1964). Dolling (1981) recognised the group as a subfamily of Cydnidae. Jacobs (1989) revised their family status and questioned their position within the Pentatomoidea, primarily because of the presence of m-chromosomes.
Contrary to the universal acceptance of the Pentatomoidea, the family-group classification is conjectural. Classifications differ in the number of accepted families, primarily because of varied ranking of certain taxa. Over the past 45 years various authors have recognised between nine and 14 families (Cassis & Gross 2002: Table 1). The classification in the Catalogue follows Schuh & Slater (1995).
Most of the above authors reject earlier notions (e.g. Kirkaldy 1909) that a number of families (Acanthosomatidae, Dinidoridae, Phloeinae, Plataspidae, Scutelleridae and Tessaratomidae) are subfamilies of the Pentatomidae. Currently, there are alternative views concerning the relationships and ranking of the Aphylidae, Cyrtocorinae, Megarididae and Thyreocoridae. The Aphylidae are either considered as a family (Schuh & Slater 1995) or a subfamily of Pentatomidae (Gross 1975; Carver et al. 1991). The Cyrtocorinae are either recognised as a pentatomid subfamily (Schuh & Slater 1995) or elevated to family (Gapud 1991; Packauskas & Schaefer 1998). Similarly, Megarididae are placed in the Pentatomidae (Gapud 1991) or recognised at family rank (Schuh & Slater (1995). The Thyreocoridae have been considered a family (Schuh & Slater 1995) or a subfamily of Cydnidae (Dolling 1991; Schuh & Slater 1995). Of these taxa, only the Aphylidae are present in Australia, and are accepted in this Catalogue as a family (see Introduction to Aphylidae for justification).
The relationships of the family-groups within the Pentatomoidea have received scant attention. A number of comparative morphology studies have been conducted (Singh-Pruthi 1925—male genitalia; Southwood 1956—eggs; Pendergrast 1957—reproductive organs; Miyamoto 1961—gut; Kumar 1957—ejaculatory canal, 1962—genitalia and salivary glands, 1964—ejaculatory reservoir, 1965, 1969; McDonald 1966—genitalia; Schaefer 1975—abdominal trichobothria; Miyamoto 1981—heart ostia; Davidová-Vilímová 1991—pygophore, and Davidová-Vilímova & Podoubský 1999—dorsal abdominal glands and androconia).
Most of the above authors drew conclusions about in-group relationships in the Pentatomoidea but in the main their data were not analysed using testable phylogenetic methods. Gapud (1991) alone provided a phylogenetic schema based on character analysis, and established the following sister-group relationships: Urostylidae + Phloeidae + (Dinidoridae + (Tessaratomidae) + Acanthosomatidae + (Scutelleridae + Canopidae) + (Cydnidae + (Lestoniidae + Plataspidae)) + Pentatomidae. Schuh and Grazia in litt. are currently testing this hypothesis.
Pentatomoids are defined by the following characters: well-developed scutellum that is either triangular to semi-elliptical in shape; claval commissure reduced or absent; paired abdominal trichobothria positioned laterally and posteriad of spiracles; base of spermatheca with a pair of ring sclerites; barrel-shaped eggs; and oval and caudally directed genital opening of male (Cobben 1968, Gapud 1991, Carver et al. 1991, Schuh & Slater 1995). Gross (1975) and Slater (1982) also give morphological diagnoses for the superfamily.
Carver, M., Gross, G.F. & Woodward, T.E. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects, etc.) [with contributions by Cassis, G., Evans, J.W., Fletcher, M.J., Hill, L., Lansbury, I., Malipatil, M.B., Monteith, G.B., Moulds, M.S., Polhemus, J.T., Slater, J.A., Štys, P., Taylor, K.L., Weir, T.A. & Williams, D.J.]. pp. 429-509 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.
Cassis, G. & Gross, G.F. 2002. Hemiptera: Heteroptera (Pentatomomorpha). in Houston, W.W.K. & Wells, A. Zoological Catalogue of Australia Vol. 27.3B. Melbourne : CSIRO Publishing, Australia. xiv 737 pp.
China, W.E. & Miller, N.C.E. 1959. Check-list and keys to the families and subfamilies of the Hemiptera-Heteroptera. Bulletin of the British Museum (Natural History) Entomology 8: 1-45
Cobben, R.H. 1968. Evolutionary Trends in Heteroptera. Part I. Eggs, architecture of the shell, gross embryology and eclosion. No. 151. Wageningen : Centre for Agricultural Publishing and Documentation Mededeling 475 pp.
Davidová-Vilímová, J. 1991. Pygophores of selected species of Pentatomoidea (Heteroptera) from Illinois. Acta Universitatis Carolinae, Biologica 35: 143-183
Davidová-Vilímová, J. & Podoubský, M. 1999. Larval and adult dorso-abdominal scent glands and androconia of central European Pentatomoidea (Heteroptera). Acta Societatis Zoologicae Bohemicae 63(1–2): 37-69
Dolling, W.R. 1981. A rationalized classification of the burrower bugs (Cydnidae). Systematic Entomology 6: 61-76
Dolling, W.R. 1991. The Hemiptera. Oxford : Oxford University Press ix 274 pp.
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). Philippine Entomologist 8(3): 865-961
Göllner-Scheiding, U. 1990. Die Entwicklung des Systems der Heteroptera (Wanzen). Entomologische Nachrichten. Berlin 34: 1-8
Gross, G.F. 1975. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera. Adelaide : A.B. James Vol. 1 250 pp., 3 colour pls.
Henry, T.J. 1997. Phylogenetic analysis of family groups within the infraorder Pentatomomorpha (Hemiptera: Heteroptera), with emphasis on the Lygaeoidea. Annals of the Entomological Society of America 90(3): 275-301
Jacobs, D.H. 1989. A new species of Thaumastella with notes on the morphology, biology and distribution of two southern African species (Heteroptera: Thaumastellidae). Journal of the Entomological Society of Southern Africa 52: 301-316
Kirkaldy, G.W. 1909. Catalogue of the Hemiptera (Heteroptera) with biological and anatomical references, lists of foodplants and parasites, etc. Prefaced by a discussion on nomenclature, and an analytical table of families. Cimicidae. Berlin : Felix L. Dames Vol. 1 xl 392 pp.
Kumar, R. 1957. On the structure and function of the so-called ejaculatory reservoir in Pentatomoidea (Hemiptera: Heteroptera). Proceedings of the Royal Society of Queensland 75: 51-65
Kumar, R. 1962. Morpho-taxonomical studies on the genitalia and salivary glands of some Pentatomoidea. Entomologisk Tidskrift 83: 44-88
Kumar, R. 1964. On the structure and function of the so called ejaculatory reservoir in Pentatomoidea (Hemiptera: Heteroptera). Proceedings of the Royal Society of Queensland 75: 51-65
Kumar, R. 1965. Contributions to the morphology and relationships of Pentatomoidea (Hemiptera: Heteroptera) Part I. Scutelleridae. Journal of the Entomological Society of Queensland 4: 41-55
Kumar, R. 1969. Morphology and relationships of the Pentatomoidea (Heteroptera) IV. Oncomerinae (Tessaratomidae). Australian Journal of Zoology 17: 553-606
Leston, D. 1958. Nomenclature of an Australasian halyine shieldbug (Hem., Pentatomidae). Entomologist's Monthly Magazine 4 19: 42
McDonald, F.J.D. 1966. The genitalia of North American Pentatomoidea. (Hemiptera: Heteroptera). Quaestiones Entomologicae 2: 7-150
Miyamoto, S. 1961. Comparative morphology of alimentary organs of Heteroptera, with the phylogenetic considerations. Sieboldia 2: 197-259
Miyamoto, S. 1981. An estimation of the gross structure of heart for the higher classification of the Heteroptera. Rostria 33: 53-66
Packauskas, R.J. & Schaefer, C.W. 1998. Revision of the Cyrtocoridae (Hemiptera: Pentatomoidea). Annals of the Entomological Society of America 91(4): 363-386
Pendergrast, J.G. 1957. Studies on the reproductive organs of the Heteroptera with a consideration of their bearing on classification. Transactions of the Royal Entomological Society of London 109: 1-63
Schaefer, C.W. 1975. Heteropteran trichobothria, (Hemiptera-Heteroptera). International Journal of Insect Morphology and Embryology 4: 193-264
Schaefer, C.W. 1993. The Pentatomomorpha (Hemiptera: Heteroptera): an annotated outline of its systematic history. European Journal of Entomology 90: 105-122
Schuh, R.T. & Slater, J.A. 1995. True Bugs of the World (Hemiptera: Heteroptera). Classification and Natural History. Ithaca : Cornell University Press xii 336 pp.
Scudder, G.G.E. 1959. The female genitalia of the Heteroptera: morphology and bearing on classification. Transactions of the Royal Entomological Society of London 111: 405-467
Singh-Pruthi, H. 1925. The morphology of the male genitalia in Rhynchota. Transactions of the Entomological Society of London 1925: 127-267
Slater, J.A. 1982. Hemiptera. pp. 417-447 in Parker, S.P. (ed.). Synopsis and Classification of Living Organisms. New York : McGraw Hill Book Co.
Southwood, T.R.E. 1956. The structure of the eggs of the terrestrial Heteroptera and its relationship to the classification of the group. Transactions of the Royal Entomological Society of London 108: 163-221
Stonedahl, G.M. & Dolling, W.R. 1991. Heteroptera identification: a reference guide, with special emphasis on economic groups. Journal of Natural History 25: 1027-1066
Štys, P. 1964. Thaumastellidae—a new family of pentatomoid Hemiptera. Acta Societatis Entomologicae Cechosloveniae 61: 236-253
Woodward, T.E., Evans, J.W. & Eastop, V.F. 1970. Hemiptera (Bugs, leafhoppers, etc.). pp. 387-457 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Carlton : Melbourne University Press 1029 pp.
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