Family MIRIDAE
Introduction
The cimicomorphan Miridae are the largest family of Heteroptera, exhibiting wide diversity in morphology, biology and host associations. The world fauna presently contains about 1300 genera and 9800 species (Schuh 1995). Henry & Wheeler (1988) estimated that the world fauna may approach 20,000 species when the rich Neotropical fauna is described. This estimate will also be increased significantly when the poorly studied Australasian and Oriental faunas are described. Mirids are a cosmopolitan group, most subfamilies having wide trans-oceanic distributions.
Mirids are mostly fragile insects, varying greatly in size, with the body elongate to ovoid. They show considerable variation in body colour, texture and vestiture. Many species are uniformly green or greenish yellow, and a number of species are brightly, possibly aposematically, coloured. The head is variable, the antennae and labium 4-segmented. Ocelli are absent, except in the Isometopinae, which have two ocelli and greatly enlarged eyes. The pronotum is pronounced and variable, often with a rounded collar (Mirinae, Deraeocorinae, Bryocorinae: Dicyphini, some Cylapinae), sometimes with callosities, and the lateral margins are sometimes constricted. The scutellum is triangular and sometimes has great expansions (Monaloniini). The hemelytra are characterised by a cuneus at the apex of the corium and one or two closed cells in the wing membrane. Wing polymophism is common with various conditions known, including aptery, coleoptery, brachyptery, microptery and macroptery. The thoracic pleura are characterised by a variable efferent system of the metathoracic scent glands, with the peritreme variously placed and shaped. The meso- and metafemora are characterised by trichobothria that are unique to the family. The pretarsus is the most important subfamilial character, and is characterised either by setiform parempodia (most Bryocorinae, Deraeocorinae, Cylapinae, Isometopinae, Phylinae) or lamellate parempodia (Orthotylinae, some Phylinae, Mirinae), and sometimes with pulvilli (Mirinae, Phylinae, Orthotylinae) or pseudopulvilli (Bryocorinae: Dicyphini, Monaloniini). The male abdomen is characterised by asymmetrical genitalia and the females have a well-developed ovipositor with complex internal genitalia (Schuh 1974–1991; Slater 1982).
Ant mimicry is very common and has evolved independently a number of times in the Miridae, with lineages in the Bryocorinae, Deraeocorinae, Mirinae, Phylinae and Orthotylinae. Morphological modifications indicating mimicry include wing shortening, constriction of the abdomen at the base, head elongation, and transverse, pale markings on the hemelytra. The most notable ant-mimetic groups include the mirine tribe Herdoniini (not Australian) and the phyline tribes Hallodapini, Leucophoropterini and Pilophorini. In Australia, the most notable ant mimics belong to the orthotyline genera Myrmecoridea Poppius and Myrmecoroides Gross, which have been placed in the Catalogue in the Orthotylinae tribe Austromirini.
The biology of mirids is reasonably well known in comparison to that of other cimicomorphans apart from the Cimicidae. The majority of species are phytophagous. They are known to feed on leaves, stems, fruits and flowers; a number of species cause severe economic damage. The genera Lygus Hahn, in North America, and Taylorilygus Leston, in Africa, are major agricultural pests. A number of phytophagous mirid pests are known from Australia, among them Creontiades dilutus (Stål), a pest of stone fruits and cotton, and the cosmopolitan potato capsid, Calocoris norwegicus (Gmelin), a serious pest of potatoes in Tasmania.
A number of mirid taxa are predaceous on eggs, larvae and adults of other arthropods, and commonly prey on small, soft bodied insects like aphids and psyllids. The Deraeocorinae and some Mirinae and Orthotylinae are beneficial natural enemies in agroecosystems. Deraeocoris Kirschbaum species are notable control agents in orchards in North America. In Australia, the phyline Tytthus mundulus (Breddin) preys on the eggs of the delphacid pest, Perkinsiella saccharicida Kirkaldy, the Sugar-cane Planthopper. The Australian endemic genus Setocoris China & Carvalho (Dicyphini) represents a special case, being cleptoparasitic on insectivorous plants of the genera Drosera L. and Byblis Salisb. Others, such as many Dicyphini species, are thought to be mixed feeders. Species of the Cylapinae are thought to be mycetophagous although their biological habits remain poorly known. Herring (1976) reported that they may feed on fungus-feeding larvae of ciid beetles in South America.
The classification of the Miridae has been based chiefly on the works of Fieber (1858) and Reuter (1910). The most comprehensive classification of the mirids derives from the catalogues and numerous papers of Carvalho (1952–1994, including co-authored papers). His work was largely followed until the phylogenetically based classifications of Schuh (1974–1991) became available. Schuh (1974, 1976) provided a major review of the suprageneric classification. He suggested major changes to the Bryocorinae, including the placement of the Dicyphini within the heterogeneous Bryocorinae. He described new tribes and reduced the Orthotylinae to a tribe of Phylinae, only to restore its subfamily status later (Schuh 1984). Schuh (1976) indicated that the tribal classifications of the Cylapinae and Deraeocorinae of Carvalho are artificial but did not provide any resolutions. Carvalho's classification remains a baseline for much of the Miridae, although many of Schuh's modifications are followed in the Catalogue. The Isometopinae and Psallopinae are regarded as valid mirid subfamilies, and the tribal classifications of the Phylinae and Bryocorinae of Schuh are followed, except for the tribal grouping of the monaloniines and dicyphines which are each ranked at tribal level. The Mirinae and Deraeocorinae are retained as separate subfamilies and the tribal classifications of these subfamilies follow Carvalho. These classifications, plus the suprageneric groups recognised in the Catalogue, are given in Table 6.
Table 6. Suprageneric classifications of Miridae. In the list by Cassis and Gross, taxa known from Australia but without described species are indicated with an asterisk, and taxa not known from Australia are given in brackets.
Carvalho (1952–1960)
Mirinae
Pithanini
Stenodemini
Mecistoscelini
Mirini
Resthenini
Hyalopeplini
Herdoniini
Orthotylinae
Halticini
Orthotylini
Pilophorini
Austromiriini
Phylinae
Phylini
Hallodapini
Dicyphini
Bryocorinae
Bryocorini
Monaloniini
Odoniellini
Deraeocorinae
Termatophylini
Deraeocorini
Saturniomirini
Cliviniemini
Hyaliodini
Surinamellini
Cylapinae
Fulviini
Cylapini
Bothriomirini
Palaucorinae
Schuh (1974–1991)
Isometopinae
Psallopinae
Phylinae
Pilophorini
Auricillocorini
Hallodapini
Leucophoropterini
Phylini
Orthotylinae
Halticini
Nichomachini
Orthotylini
Cylapinae
Mirinae
Mirini
Deraeocorini
Bryocorinae
Eccritotarsini
Dicyphini
Bryocorini
Schuh (1974–1991)
Isometopinae
Psallopinae
Phylinae
Pilophorini
Auricillocorini
Hallodapini
Leucophoropterini
Phylini
Orthotylinae
Halticini
Nichomachini
Orthotylini
Cylapinae
Mirinae
Mirini
Deraeocorini
Bryocorinae
Eccritotarsini
Dicyphini
Bryocorini
Presently, seven subfamilies, 18 tribes, 91 genera and 186 described species are recognised in the Australian mirid fauna. The fauna is actually much larger and is being revised by one of us (G.C.). Twenty-eight genera previously unrecorded from Australia are now known to have Australian representatives and numerous undescribed endemic genera are also known. Carvalho & Silva Alfonso (1977, Mirinae: Stenodemini), Carvalho & Lorenzato (1978, Cylapinae), Carvalho & Gross (1980, Mirinae: Hyalopeplini), Carvalho (1981, Bryocorinae: Monaloniini), Schuh (1984, Phylinae) and Stonedahl (1988, Bryocorinae: Eccritotarsini) have provided generic and species reviews of the Indo-Pacific fauna which are relevant to the Australian fauna.
The Isometopinae are considered to be the most plesiomorphic subfamily of Miridae. In common with the rest of the Miridae, they have femoral trichobothria, asymmetrical male genitalia and sclerotised rings in the female genitalia. Eyles (1971, 1972) provided checklists of the world species and Ghauri & Ghauri (1983) provided a generic key to the world fauna. Henry (1980, 1984) provided a key to the Western Hemisphere genera and a revision of the North American species. Herczek (1993) revised the systematic position of the Isometopinae, reviewed their morphology, and proposed a new suprageneric classification. His tribal classification is not used in the Catalogue. The Australian fauna contains the endemic Eurocrypha thanatochlamys Kirkaldy and a number of undescribed taxa.
The Psallopinae are a small subfamily, probably most closely related to the Isometopinae and found in the Indo-Pacific and South Africa. An undescribed species of Psallops usinger is known from Queensland.
The Cylapinae are recognised by a simple pretarsus and a rounded collar and are grouped with the Psallopinae and Isometopinae on the basis of the presence of a subapical tooth on the tarsal claws (Schuh & Schwartz 1984). The Cyalpini are represented by two coleopteroid genera, Carvalhoma Slater & Gross and Schizopteromiris Schuh, with the latter genus found in Queensland, Lord Howe Island and New Caledonia (Schuh 1986). The Fulviini are presently represented by five genera, each containing one species. Numerous new species of Cylapofulvius Poppius and Fulvius Stål require description. The genus Peritropis Uhler, previously not recorded from Australia, contains numerous undescribed species.
The Phylinae are one of the largest subfamilies. Most species are 2 to 5 mm in length. Schuh (1984) redefined the Phylinae as a monophyletic group on the basis of the male genitalia, with the phallotheca attached to the posterior wall of the genital capsule. The Phylini are possibly paraphyletic and are represented in Australia by a largely undescribed fauna. Schuh's (1984) monograph of the Indo-Pacific Phylinae is a relevant generic framework for the Australian fauna and also indicates that undescribed species of Gulacapsus Schuh and Arafuramiris Schuh occur in Australia. Malipatil (1992) reviewed the Australian Campylomma Reuter fauna, describing new species and providing new host plant information. Carvalho & Gross (1982) revised the Australian Leucophoropterini, recognising seven genera and 42 species. The ant-mimetic tribe Hallodapini contains two genera, each with one species, but numerous new genera and species await description.
The Orthotylinae are defined by convergent lamellate parempodia and distinctive male genitalia. Three tribes are represented in the Australian fauna: the endemic Austromirini including the genera Austromiris Kirkaldy, Dasymiris Poppius, Pophyrodema Reuter and Zanessa Kirkaldy; Orthotylini with nine genera and 11 species, including widespread genera such as Orthotylus Fieber and Melanotrichus Reuter represented by only a few species; and Halticini with two species of Halticus Hahn, including the pioneer species, H. minutus Reuter, and the distinctive genus Coridromius Signoret.
The Mirinae are represented by three tribes which share the feature of divergent lamellate parempodia. The tribe Mirini contains 16 genera and 32 species in Australia. Together with the Phylinae, Mirinae are probably the most speciose subfamily, with many new taxa to be described. The genera Taylorilygus and Creontiades Distant contain the majority of species, with most other genera containing only one species. The grass-inhabiting tribe Stenodemini is depauperate in Australia and contains only five species. The clear-winged Hyalopeplini contains five genera and 11 species. The mirine species, Paralygus simplex Reuter, 1905, was described from Sydney, Australia. Carvalho (1976) indicated that this species is a synonym of a Mexican species, Proba inspersa (Distant, 1884), and that the type specimen was mislabelled. This interpretation has been accepted and this species has not been included in the Catalogue.
Three of the six tribes of the Deraeocorinae recognised by Carvalho (1955) are found in Australia. The Deraeocorini from Australia are robust, mostly punctate mirids with setiform parempodia and no pulvilli; the tribe comprises six genera and eight species. The Australian species of Fingulus Distant was recently redescribed by Stonedahl & Cassis (1991) in a revision of this Indo-Pacific genus. The Australasian tribe, the Saturniomirini, is represented in Australia by two genera and six species. Cassis (1995) revised the Termatophylini of the world and recognised three genera and six species from Australia. He reviewed their biology and recorded two species, Kundakimuka queenslandica Cassis and Termatophylum melaleucae Cassis from Melaleuca species, the former species feeding on lepidopteran larvae.
The subfamily Bryocorinae is a heterogeneous group of four tribes. The Monaloniini contains species which often have expansions of the scutellum. They are related to the Dicyphini on the basis of the pretarsal structure and contain, outside Australia, a number of species that are major pests of crops such as cocoa, tea and coffee (Stonedahl 1991). Commonly encountered species include Felisacus elegantulus (Reuter) the fern-inhabiting species, and Eucerocoris suspectus Distant which feeds on Melaleuca (Cav.) S.T.Blake [Myrtaceae] and is presently considered as a potential biological control agent of Melaleuca quinquenerva (Cav.) S.T.Blake, a major weed in south-eastern United States. The Dicyphini contains four genera, all of which inhabit sticky plants. Many new species of Setocoris require description. The Eccritotarsini is represented in Australia by undescribed species and the Bryocorini contains two species.
General References
Carvalho, J.C.M. 1952. On the major classification of the Miridae (Hemiptera). (With keys to subfamilies and tribes and a catalogue of the world genera). Anais da Academia Brasileira de Ciências 24: 31-110
Carvalho, J.C.M. 1955. Keys to the genera of Miridae of the World (Hemiptera). Boletim do Museu Paraense Emilio Goeldi 11(2): 5-151
Carvalho, J.C.M. 1957. Catálogo dos Mirídeos do Mundo. Parte I. Subfamílias Cylapinae, Deraeocorinae, Bryocorinae. Arquivos do Museu Nacional. Rio de Janeiro 44: 1-158
Carvalho, J.C.M. 1958. Catálogo dos Mirídeos do Mundo. Parte II. Subfamília Phylinae. Arquivos do Museu Nacional. Rio de Janeiro 45: 1-216
Carvalho, J.C.M. 1958. Catálogo dos Mirídeos do Mundo. Parte III. Subfamília Orthotylinae. Arquivos do Museu Nacional. Rio de Janeiro 47: 1-161
Carvalho, J.C.M. 1959. Catálogo dos Mirídeos do Mundo. Parte IV. Subfamília Mirinae. Arquivos do Museu Nacional. Rio de Janeiro 48: 1-384
Carvalho, J.C.M. 1960. Catálogo dos Mirídeos do Mundo. Parte V. Bibliografia e Indice Geral. Arquivos do Museu Nacional. Rio de Janeiro 51: 1-194
Carvalho, J.C.M. 1976. Analecta Miridologica: concerning changes of taxonomic position of some genera and species (Hemiptera). Revista Brasileira de Biologia 36: 49-59
Carvalho, J.C.M. 1981. The Bryocorinae of Papua New Guinea (Hemiptera, Miridae). Arquivos do Museu Nacional. Rio de Janeiro 56: 35-89
Carvalho, J.C.M. & Gross, G.F. 1979. The tribe Hyalopeplini of the World (Hemiptera: Miridae). Records of the South Australian Museum 17: 429-531
Carvalho, J.C.M. & Gross, G.F. 1982. Australian ant-mimetic Miridae (Hemiptera: Heteroptera). I. The Leucophoroptera group of the subfamily Phylinae. Australian Journal of Zoology Supplementary Series 86: 1-75
Carvalho, J.C.M. & Lorenzato, L.M. 1978. The Cylapinae of Papua New Guinea (Hemiptera: Miridae). Revista Brasileira de Biologia 38: 121-149
Carvalho, J.C.M. & Silva Afonso, C.R. da 1977. The tribes Mecistoscelini and Stenodemini in Papua-New Guinea (Hemiptera: Miridae). Revista Brasileira de Biologia 37: 807-823
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. 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. Proceedings of the Entomological Society of Washington 97: 258-330
Eyles, A.C. 1971. List of Isometopidae (Heteroptera: Cimicoidea). New Zealand Journal of Science 14: 940-944
Eyles, A.C. 1972. Supplement to list of Isometopidae (Heteroptera: Cimicoidea). New Zealand Journal of Science 15: 463-464
Fieber, F.X. 1858. Criterien zur generischen Theilung der Phytocoriden (Capsini aut.). Wiener Entomologische Monatschrift 2: 289-327
Ghauri, M.S.K. & Ghauri, F.Y.K. 1983. A new genus and new species of Isometopidae from north India, with a key to the world genera (Heteroptera). Reichenbachia 21: 19-25
Henry, T.J. 1980. Review of Lidopus Gibson and Wetmorea McAtee & Malloch, descriptions of three new genera and two new species, and key to New World genera (Hemiptera: Miridae: Isometopidae). Proceedings of the Entomological Society of Washington 82: 178-194
Henry, T.J. 1984. New species of Isometopidae (Hemiptera: Miridae) from Mexico, with new records for previously described North American species. Proceedings of the Entomological Society of Washington 86: 337-345
Henry, T.J. & Wheeler, A.G., Jr 1988. Miridae. pp. 251-507 in Henry, T.J. & Froeschner, R.C. (eds). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental United States. Leiden : E.J. Brill xix 958 pp.
Herczek, A. 1993. Systematic Position of Isometopinae Fieb. (Miridae, Heteroptera) and their Intrarelationships. Katowice : Uniwersytet Šlaski 86 pp.
Herring, J.L. 1976. A new genus and species of Cylapinae from Panamá (Hemiptera: Miridae). Proceedings of the Entomological Society of Washington 78: 91-94
Malipatil, M.B. 1992. Revision of Australian Campylomma Reuter (Hemiptera: Miridae: Phylinae). Journal of the Australian Entomological Society 31: 357-368
Reuter, O.M. 1910. Neue Beiträge zur Phylogenie und Systematik der Miriden nebst einleitenden Bemerkungen über die Phylogenie der Heteropteren-Familien. Acta Societatis Scientiarum Fennicae 37(3): 1-167
Schuh, R.T. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with a phylogenetic analysis of the ant-mimetic tribes of the two subfamilies for the world. Entomologica Americana 47: 1-332
Schuh, R.T. 1975. The structure, distribution, and taxonomic importance of trichobothria in the Miridae (Hemiptera). American Museum Novitates 1975(2585): 1-26
Schuh, R.T. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of relationships within the family. American Museum Novitates 1976(2601): 1-39
Schuh, R.T. 1984. Revision of the Phylinae (Hemiptera, Miridae) of the Indo-Pacific. Bulletin of the American Museum of Natural History 177: 1-476
Schuh, R.T. 1986. Schizopteromiris, a new genus and four new species of coleopteroid Cylapinae Miridae from the Australian region (Heteroptera). Annales de la Société Entomologique de France ns 22: 241-246
Schuh, R.T. 1991. Phylogenetic, host and biogeographic analyses of the Pilophorini (Heteroptera: Miridae: Phylinae). Cladistics 7: 157-189
Schuh, R.T. 1995. Plant Bugs of the World (Heteroptera: Miridae). Systematic Catalog, Distributions, Host List and Bibliography. New York : New York Entomological Society 1329 pp.
Schuh, R.T. & Schwartz, M.D. 1984. Carvalhoma (Hemiptera: Miridae); revised subfamily placement. Journal of the New York Entomological Society 92: 48-52
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
Stonedahl, G.M. 1988. Revisions of Dioclerus, Harpedona, Mertila, Myiocapsus, Prodromus, and Thaumastomiris (Heteroptera: Miridae: Bryocorinae: Eccritotarsinae). Bulletin of the American Museum of Natural History 187: 1-99
Stonedahl, G.M. 1991. The Oriental species of Helopeltis (Heteroptera: Miridae): A review of economic literature and guide to identification. Bulletin of Entomological Research 81: 465-490
Stonedahl, G.M. & Cassis, G. 1991. Revision and cladistic analysis of the plant bug genus Fingulus Distant (Heteroptera: Miridae: Deraeocorinae). American Museum Novitates 3038: 1-55