Compiler and date details
15 February 2002
The Pentatomidae or stinkbugs are the most diverse family of pentatomomorphan bugs, numbering about 900 genera and 4700 species (Schuh & Slater 1995; Henry 2009; Cassis & Gross 2002: Table 12). They are found in all major zoogeographic regions and are particularly diverse in the subtropical and tropical regions of the world. The family is represented in Australia by 134 genera and 363 species, of which 94 genera and 330 species are endemic.
The taxonomic arrangement presented by Cassis & Gross (2002) was greatly influenced by an unpublished work by Dr David Rider (in litt.). His work in preparation on the tribal classification for the Pentatominae includes the placement of all genera and was followed by Cassis & Gross (2002) so that the Australian Pentatomidae fauna can be placed within a global context. Much of this arrangement was the original work of Dr Rider which was acknowledged.
The classification of the Pentatomidae remains conjectural (Schuh 1986; Schuh & Slater 1995) and requires urgent attention. The ranking of a number of taxa (e.g. Aphylidae, Scutelleridae) varies between pentatomoid family and pentatomid subfamily in the alternative classifications of workers (Cassis & Gross 2002: Table 13, also General and Pentatomoidea introductions). Leach (1815) first accorded family status to the group. Stål (1865, 1866, 1868, 1870, 1876) established the basis of the modern classification. Kirkaldy (1909) catalogued the world fauna.
The subfamilial classification has varied in composition and division. From Kirkaldy (1909) to the present, the number of subfamilies recognised has varied from two to thirteen subfamilies (Cassis & Gross 2002: Table 13). Earlier authors (e.g. Kirkaldy 1909; Miller 1956; China & Miller 1959) had a broader concept of the family, incorporating the dinidorids, plataspids, tessaratomids and scutellerids. These latter taxa are now consistently given individual family status. Gross (1975a) rejected most previous classifications, because of regional biases and lack of morphological support. He established two subfamilies: the Aphylinae, a monogeneric subfamily, and the remainder of pentatomids in a composite concept of the Pentatominae. Within the Pentatominae he erected 27 informal 'genus-groups', diagnosed on the basis of external and genitalic characters. Carver et al. (1991) followed this arrangement. Gapud (1991) in a review of pentatomoid and pentatomid relationships recognised nine subfamilies. This classification received partial support from some authors. Schuh & Slater (1995) recognised family status for the Aphylidae and Megarididae, and subfamily for the Edessinae (also Rolston & McDonald 1979). Rider (in litt.) differs from Schuh & Slater (1995) only in the ranking of the aphylids, accepting them as a subfamily of the Pentatomidae. Rider (2006) provided a Catalogue to the Palaearctic Pentatomidae. Konstantinov & Gapon (2005) discussed the structures of the male genitalia in the subfamilies Discocephalinae and Phyllocephalinae.
Four of the eight pentatomid subfamilies (sensu Rider in litt.) are represented in Australia: the Asopinae, Pentatominae, Phyllocephalinae and Podopinae. The Australian fauna represents 10% of the world pentatomid genera and about 8% of the species. The endemicity of the fauna is notable, with 70% of the genera and over 90% of the species restricted to Australia (Table 12).
The majority of subfamilies that are not represented in Australia are found in the Western Hemisphere. The Cyrtocorinae are a small subfamily (four genera and 11 species) confined to the Neotropical Region. The Discocephalinae comprise around 70 genera and more tham 260 species and are most diverse in the neotropics.
The Edessinae are found in the Western Hemisphere, predominantly in South America, and comprise four genera and around 270 species. Rolston & McDonald (1979) and Schuh & Slater (1995) provided reviews of these extralimital taxa.
The Serbaninae are a monobasic group restricted to Kalimantan (Leston 1953).
The Asopinae are a cosmopolitan subfamily of predatory bugs, comprising around 65 genera and more than 350 species (Schuh & Slater 1995). Thomas (1992, 1994) revised the Asopinae of the world and established that two-thirds of the genera and half of the species occur in the Eastern Hemisphere. The asopines are depauperate in Australia, represented by only eight genera (1 endemic) and 11 species (3 endemic). Australian species that also occur extralimitally are often found in Melanesia and/or the Oriental Region; Cermatulus nasalis (Westwood) is also found in New Zealand. The asopine fauna of Australia is most often represented in collections by the ubiquitous Oechalia schellenbergii (Guérin) which is found in all States and territories of mainland Australia. Thomas (1994) diagnosed all the Australian taxa.
The Phyllocephalinae are restricted to the Eastern Hemisphere (chiefly Indo-Pacific) and comprise around 30 genera and more than 170 species. Ahmad & Kamaluddin (1988, 1990) and Kamaluddin & Ahmad (1988) established a tribal classification, recognising four tribes: Cressonini, Megarrhamphini, Phyllocephalini and Tetrodini. The nominotypical tribe is the only tribe represented in Australia and comprises three genera (2 endemic) and six species (5 endemic). All the species are broadly distributed in Australia, often in more xeric climes. Gross (1975a) suggested that the Phyllocephala genus-group is related to the Cephaloplatus genus-group. Gross (1976) revised the South Australian fauna, describing all the genera and five of the species.
The Podopinae are a cosmopolitan subfamily comprising around 65 genera and more than 250 species worldwide (Schuh & Slater 1995). Davidová-Vilímová & Štys (1993) and Davidová-Vilímová & McPherson (1995) proposed a new tribal classification as follows: Brachycerocorini, Deroploini, Graphosomatini, Podopini and Tarisini. This arrangement is followed in the Catalogue. The Australian fauna is represented by two tribes (Deroploini and Podopini), 11 genera and 22 species. The Deroploini is almost exclusively Australian with eight genera and 19 species restricted to Australia; Protestrica subpunctata (Walker) is known from Papua New Guinea and is the only species of the tribe found outside Australia. The Podopini is the most diverse tribe of the Podopinae, with 27 described genera. The tribe is only represented in Australia by two species, Coracanthella geophila (Montrouzier) and Scotiniphora allanae (Musgrave), both of which occur primarily in northern Australia and also extralimitally in the Eastern Hemisphere. Musgrave (1930) described the majority of the Australian podopine species, describing two new genera and eight new species. Gross (1975a) recognised two generic-groups for the Podopinae: Podops genus-group and Tarisa genus-group; he suggested that the two groups are polyphyletic. He also described the Deroploini from South Australia, with five genera and 11 species.
The Pentatominae are one of the most diverse subfamilies of heteropteran bugs and comprise around 400 genera and more than 2770 species (Schuh & Slater 1995). The tribal classification of the Pentatominae is in a state of significant confusion and different taxonomic arrangements have been proposed (Table 14). Many attempts to divide the subfamily are characterised by a number of small to medium-sized tribes, with the majority of taxa included in the nominotypical tribe. Kirkaldy (1909) divided the subfamily into ten tribes with over half the genera and species assigned to the Pentatomini. Gross (1975a) in a critique of previous suprageneric classifications of the subfamily noted that most alternatives were largely regional in scope. He addressed this shortfall by erecting 27 'genus-groups' to accommodate the Australian fauna and provided a schema of relationships. Hasan & Kitching (1993) provided a cladistic analysis of various taxa to resolve the fragmentary nature of previous classifications. Their work is confounded by a priori acceptance of the monophyly of most of the terminal taxa (aside from the Podopini and Strachini). Unfortunately, they did not present a revised classification based on their analysis; the tribes in Table 14 are a mere listing of their study taxa.
Rider's (in litt.) catalogue of the world fauna is the first of its kind since Kirkaldy (1909). This work lists 42 tribes on a worldwide basis and includes the placement of nearly all described genera within this classification. Nineteen of these tribes are represented in Australia (Table 14) and the majority of genera and species are assigned to these groups. Twenty-one genera and 45 species remain unplaced in the Catalogue.
The Aeptini is a small tribe found in the Afrotropical and Australian Regions. Stål (1871) and Kirkaldy (1909) assigned Australian genera to this tribe, although Gross (1975a) was sceptical of this arrangement. This arrangement is maintained in the Catalogue, the tribe being represented by seven genera and 18 species, all of which are endemic. These taxa are found across Australia, often in more xeric climes. Gross (1975a) revised the South Australian fauna (= as Menestheus genus-group) and provided habitus and genitalic illustrations.
The Agonoscelidini is a monogeneric tribe that is restricted to the Eastern Hemisphere. Its greatest diversity occurs in the Afrotropical and Oriental Regions. The Australian fauna is represented by the Horehound Bug, Agonoscelis rutila (Fabricius), which is found broadly in eastern Australia, as well as the Torres Strait Islands, and in Melanesia and South-East Asia (Malaysia as the northern limit).
The Antestini is a diverse tribe of pentatomines, possibly confined to the Eastern Hemisphere. The tribe is represented in Australia by four genera and 10 species, of which one genus and eight species are endemic. The majority of species occur in northern Australia, and two species of Plautia Stål are also found in Melanesia. Gross (1975a) established the Antestia genus-group, which, in addition to the nominotypical genus, included Anaxilaus Stål, Antestiopsis Leston and Novatilla Distant, but he removed Plautia to his Pentatoma genus-group. In the Catalogue we follow the arrangement of Rider (in litt.), with Plautia retained in the tribe.
The Axiagastini is a small tribe that is restricted to South-East Asia, Melanesia and Australia. The Australian fauna includes a single species, Axiagastus rosmarus Dallas, found in tropical north Queensland, Papua New Guinea, Indonesia, Malaysia and the Philippines.
The Carpocorini is one of the most diverse tribes of Pentatominae and is cosmopolitan in distribution. The tribe is represented in Australia by seven genera and eleven species, of which five genera and ten species are endemic. Many of the species are broadly distributed in the temperate regions of Australia, with Monteithiella humeralis Gross also found in New Zealand. Gross (1976) established the Carpocoris genus-group for the inclusion of these genera, as well as Agonoscelis Spinola and Platycoris Guérin as (= Hypogomphus Spinola). The latter two genera are placed elsewhere in the Catalogue (Agonoscelidini and Haylini respectively). Mycoolona Distant is included in the Carpocorini based on the classification of Rider (in litt.). Aside from Gross' (1976) review of the South Australian fauna there has been no modern treatment of the Australian fauna.
The Catacanthini is a small tribe that also has an Indo-Pacific distribution (including Madagascar). The genus Catacanthus Spinola is represented in Australia by a single species currently known from Cape York and Java. Gross (1975a), Ahmad & Kamaluddin (1981) and Monteith (1993) dealt with the systematics and biology of this species.
The Caystrini is primarily an Indo-Pacific tribe (Ahmad & Afzal 1979) and is represented in Australia by two species that are found in tropical north Queensland and extralimitally in Papua New Guinea and the Oriental Region.
The Diemeniini is restricted to the Australian Region (Australia and Papua New Guinea) and comprises 13 genera and 49 species. In the Catalogue the tribe is more limited in composition than proposed by Kirkaldy (1909) who included the Afrotropical genus Mecidea Dallas (now in Mecideini). Gross (1976) established the Diemenia genus-group for the Australian representatives and described many of the taxa in his review of the South Australian fauna. In Australia the tribe is represented by 13 genera with all but one (Oncocoris Mayr) restricted to Australia. The most diverse genera are Diemenia Spinola (4 species), Kalkadoona Distant (6 species), Niarius Stål (6 species) and Oncocoris Mayr (20 species). Many of the species are broadly distributed in the arid and semi-arid regions of Australia. There appear to be both Bassian and Eyrean components of the diememiine fauna. McDonald & Edwards (1978) revised Oncocoris and described eight new species. The assignment of Caridophthalmus Assman to this tribe is conjectural. Ahmad et al. (1984) revised the genus Aplerotus Dallas, which is known to occur in Australia, as well as Melanesia and Indonesia.
The Eysarcorini (= Eysarcoris genus-group sensu Gross 1976) is a moderately diverse tribe that is confined to the Eastern Hemisphere. The tribe is represented in Australia by two genera (Eysarcoris Hahn and Spermatodes Bergroth) and eight species, seven of which are endemic. The species are mostly distributed in the tropics of Australia; Wood & McDonald (1984) and McDonald (1989) reviewed the Australian fauna.
The Halyini is a cosmopolitan tribe that is extremely diverse, particularly in the Afrotropical and Australian Regions. Gross (1976) established the Halys genus-group, suggesting that it is primarily an Eastern Hemisphere group, and dividing the fauna of the Australian region into three sections. The tribe is the most diverse suprageneric group of the Pentatomidae found in Australia and is represented by 19 genera and 80 species (70 endemic species). Fourteen of the genera are Australian endemics, with Anchises Stål, Austromalaya Kirkaldy, Bromocoris Horváth and Poecilometis Dallas also found in Papua New Guinea. The Australian fauna is dominated by Poecilometis, which comprises 38 species and is found across the continent. Gross (1972) revised the genus and described 12 new species and three new subspecies. He subsequently (Gross 1976) revised the South Australian fauna and provided a key to ten genera. McDonald (1992) reviewed the genus Munduala Distant and described two new species. McDonald (1995) also revised the genus Platycoris Guérin (as Hypogomphus Spinola).
The Menidini (= Menida genus-group of Gross 1976) is a moderately diverse tribe that is primarily found in the Eastern Hemisphere. The tribe is represented in Australia by four genera (1 endemic) and 10 species (6 endemic). The nominotypical genus, Menida Motschulsky, is represented in Australia by six species. Most of the species of the tribe are distributed in the tropical north of Australia. The Australian taxa have received little taxonomic attention aside from the regional treatment of Gross (1976).
The Myrocheini is an Eastern Hemisphere tribe that is most diverse in the Oriental Region and encompasses the Dictyotus and Tholosanus genus-groups of Gross (1975a, 1976). In Australia, the tribe is represented by five genera and 14 species, of which four genera and 10 species are endemic. Dictyotus is the most speciose genus, comprising nine species, a number of which are broadly distributed in the temperate zone of Australia. Gross (1975a, 1976) revised the South Australian fauna and reviewed the complex nomenclature of Dictyotus.
The Nezarini is a moderately diverse group that is cosmopolitan in distribution, but is indigenous in the Eastern Hemisphere. The group is most notable for the green vegetable bug (or southern green stink bug), Nezara viridula (Linnaeus), which is a serious pest in many parts of the world. Clarke (1992) reviewed the distribution of this species in Australia, where it is found mainly in the east. The tribe is also represented in Australia by Alciphron Stål (1 species) and Glaucias Kirkaldy (2 species), all of which are found extralimitally in Melanesia and/or Oriental Region.
The Pentatomini is cosmopolitan in distribution and is the most diverse tribe of the subfamily. The tribe is depauperate in Australia being represented by only two genera (Amblycara Bergroth and Hyrmine Stål) and three species.
The Piezodorini is a circumtropical tribe that includes only the single species of the nominotypical genus, Piezodorus Fieber. Until recently, the Asian soybean pest species, P. hybneri Gemlin, was thought to occur in Australia. Staddon & Ahmad (1995) and Staddon (1997) claimed that this was a misidentification. Both Ahmad (1995) and Staddon (1997) described a new species for the Australian and Pacific populations. Unfortunately, they separately established available names with the same species epithet, P. grossi Ahmad and P. grossi Staddon, for this taxon. We are proposing that another existing name is available for this species, P. oceanicus (Montrouzier), and establish a new synonymy, that includes the aforementioned available names.
The Rhynchocorini (= Rhynchocoris genus-group sensu Gross 1976) is a diverse tribe of pentatomines restricted to the Oriental and Australian Regions, and most diverse in Australia. The tribe is represented on the continent by 14 genera and 57 species, of which nine genera and 53 species are endemic. The most diverse genera are Cuspicona Dallas (21 species), Diaphyta Bergroth (7 species), Ocirrhoe Stål (11 species) and Vitellus Stål (7 species). Many of the species of the tribe are distributed in the tropical and subtropical regions of Australia. Gross (1975b, 1976) revised the Australian fauna, providing a key to genera and recognising 38 species. Gross & McDonald (1994) revised Avicenna Distant and Vitellus and described new species. McDonald (2001) reviewed the genus Diaphyta.
The Sciocorini is primarily native to the Eastern Hemisphere, with the majority of species occurring in the Palaearctic Region (Kirkaldy 1909). The tribe is represented in the drier parts of Australia by three genera and four endemic species. Gross (1976) grouped these taxa in his Kapunda genus-group, and described the two known Kapunda Distant species.
The Strachini (= Strachia genus-group sensu Gross 1976) is cosmopolitan in distribution and is most diverse in the Eastern Hemisphere. The group is relatively depauperate in Australia, being represented by only two genera and four endemic species. Ahmad & Khan (1980, 1983) described an endemic monotypic genus, Grossiana Ahmad & Khan, and reviewed the taxonomy and zoogeography of Stenozygum Fieber.
A large number of taxa (21 genera and 45 species) remained unplaced in Cassis & Gross (2002). Aside from one genus and species, all are endemic to Australia. The most notable genus is Cephaloplatus White which contains 17 species (Gross 1970). Cassis & Gross (2002) refer to the position of these unplaced taxa in Gross' (1975a, 1976) genus-group classification.
The biology of Pentatomidae is comparatively well-known and useful summaries include those of Gross (1975a, 1976), Dolling (1991) and Schuh & Slater (1995). Most pentatomids are phytophagous and feed on the plant vascular system (leaves and stems) or immature fruits and seeds (Schuh & Slater 1995). Panizzi et al. (2000) provided a detailed overview of the species of economic importance. They reported that pentatomid feeding causes wilting, abortion of fruit and seeds, and in some cases stinkbugs transmit plant pathogens. The worldwide pest species, Nezara viridula, is also a serious pest in Australia, and causes damage to foodcrops, including sorghum, soybean, lucerne, sunflower, cotton, potatoes and numerous garden vegetables (Hely et al. 1982). Clarke (1992) summarised the distribution and pest status of this species. The Spined Citrus Bug, Biprorulus bibax Breddin, is a serious pest of citrus; its biology was summarised by Panizzi et al. (2000). Other species that have occasional economic significance include: Dictyotus caenosus (Westwood) on soybeans, Vitellus Stål species on citrus, Plautia afffinis Dallas on a variety of fruits and vegetables, Cuspicona simplex Walker on potatoes and tomatoes, Agonoscelis rutlia (Fabricius), an occasional pest on grain legumes and fruit trees, and Piezodorus oceanicus on cucurbits.
The phytophagous Penatomidae of Australia are known to occur on a broad range of host plant families. Cassis & Gross (2002) recorded them from 37 plant families across the Angiospermae and 234 host species records (see Cassis & Gross 2002: Tables 5, 6 and 7 General Introduction and Appendix IV). They are most frequently associated with the rosid eudicots, particularly the Myrtaceae, Fabaceae and Mimosaceae. The Pentatominae: Halyinae are commonly associated with myrtaceous plants, particularly species of the genus Eucalyptus l'Hér. Members of the Diemeniini are also associated with eucalypts, and genera such as Diemenia Spinola and Niarius Stål are found under bark. Several pentatomids are known to feed on grasses and evidence is accumulating that members of the Aeptini may be specialised grass-feeders.
Asopinae are primarily predacious. Little is known of the biology of the Australian species. Andrallus spinidens (Fabricius), Cermatulus nasalis (Westwood) and Oechalia schellenbergi (Guérin) are known to feed on larvae of Lepidoptera.
Pentatomids are mostly ovoid in shape, although a few species are elongate. The body is dorsally flattened and ventrally convex. The antennae are usually 5-segmented, although occasionally 3 or 4-segmented. The pronotum is laterally angulate. The scutellum is usually triangular and large, with a few species having the scutellum greatly enlarged and shield-like. The hemelytra are well developed, with 5–12 membrane veins. The hind wings do not have a hamus. The tarsi are 2 or 3-segmented. The trichobothria are arranged transversely caudal to the spiracles. The ovipositor is laciniate. The aedeagus has one to three pairs of conjunctival appendages. (Slater 1982; Schuh & Slater 1995)
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