Infraorder PENTATOMOMORPHA
Compiler and date details
June 2012 - Professor Gerry Cassis, Anna Namyatova, Nikolai Tatarnic and Celia Symonds, University of New South Wales, Sydney
15 February 2002 - Gerasimos Cassis, Australian Museum, Sydney, New South Wales, Australia & Gordon Gross, South Australian Museum, Adelaide, South Australia, Australia
Introduction
[Introduction after Cassis & Gross (2002), with minor updating of numbers of described taxa recorded for Australia].
The Heteroptera is the most biologically and taxonomically diverse suborder of Hemiptera, a paraneopteran order of exopterygote insects. The suborder has been divided into seven infraorders: Enicocephalomorpha, Dipsocoromorpha, Gerromorpha, Leptopodomorpha, Nepomorpha, Cimicomorpha and Pentatomomorpha (Štys & Kerzhner 1975; Schuh 1979; Carver et al. 1991; Schuh & Slater 1995). This present list is based on the Zoological Catalogue of Australia Volume, 27.3B, now updated here to include 34 families, 449 genera, 1203 species for the Australian Pentatomomorpha. The Australian taxa of all other infraorders were catalogued in the Zoological Catalogue of Australia Volume, 27.3A (Cassis & Gross 2002), also now updated in the Australian Faunal Directory (AFD).
The Pentatomomorpha constitutes the most recognisable group of true bugs, and includes the flat bugs, burrower bugs, stink bugs, shield bugs, chinch bugs, stilt bugs, pod bugs and cotton stainers. Unlike the other infraorders, in which carnivory is a common feeding behaviour, the pentatomomorphan bugs are chiefly phytophagous. Omnivory and carnivory (e.g. in Pentatomoidea: Asopinae and Geocoridae) are less common modes of feeding. The Pentatomomorpha are generally found in association with eudicotyledonous angiosperms or monocots (Blissidae, Colobathristidae, Pachygronthidae). There are also specialised bark-dwelling (Aradidae) and epigaeic groups (e.g. Rhyparochromidae), and termite inquilines (Termitaphididae).
In many families of the Lygaeoidea and Coreoidea, granivory is the generalised feeding habit, whereas in the Pentatomoidea feeding on plant vessels is the norm. As in the cimicomorphan superfamily Miroidea, monophagy occurs, such that inferences can be made about host associations. This infraorder also contains several economically important species, most of which are pests in agro-ecosystems.
All published genus and species synonymies, including extralimital synonymies are given in the AFD. Where classifications are in dispute, alternative views are listed. All relevant literature from 1758 to May 2012 is included in this current checklist. The family introductions provide a current review of the world fauna, including an account of their classification, diversity, morphology and biology, as well as a synopsis of the Australian fauna. For general accounts of the classification and biology of the Heteroptera, users are referred to Carver et al. (1991), Dolling (1991), Schuh & Slater (1995) and Schaefer & Panizzi (2000).
CLASSIFICATION
Alternative classifications of the Pentatomomorpha have been proposed recently that vary in the number of superfamilies and families, and the composition of each superfamily. The number of families recognised ranges from 29 to 40, and there are differences in the acceptance of the Idiostoloidea as a superfamily. A sample of the most cited classifications is listed in Table 1 Cassis & Gross (2002).
Much of the classification in Cassis & Gross (1995) followed the synthetic classifications of Schuh & Slater (1995) and Henry (1997). These classifications are based on phylogenetic analyses (see also Schuh 1979, 1986) and are the most rigorously derived and testable classifications currently available. The classification used in Cassis & Gross (2002) differed markedly from both that of Carver et al. (1991) and the classification we proposed in an earlier work (Cassis & Gross 1995—Table 2). Departures from these latter classifications are necessary to align with modern developments in Heteropterology.
All of the pentatomomorphan superfamilies and 34 of the families are represented in Australia. The Urostylidae, previously recognised in Australia (Woodward et al. 1970; Carver et al. 1991), were removed from the fauna.
Infraordinal Classification
Leston et al. (1954) established the basis of the modern classification of the Heteroptera, and in doing so, proposed the Aradidae + 'Trichophora' as the fundamental component of the Pentatomomorpha. Štys & Kerzhner (1975) transferred the Leptopodidae, Saldidae and Thaumastocoridae from the Pentatomomorpha, and narrowed the concept of the infraorder. The monophyly of the Pentatomomorpha has been upheld by most subsequent authors (Štys & Kerzhner 1975; Cobben 1978; Schuh 1986; Carver et al. 1991; Wheeler et al. 1993; Schuh & Slater 1995).
Schaefer (1975, 1993) supported an earlier grouping, the 'Trichophora' (Tullgren 1918), for those pentatomomorphan taxa that possess patterned abdominal trichobothria. Henry (1997) established this taxon (Lygaeoidea + Coreoidea + Pyrrhocoroidea + Pentatomoidea) as a monophyletic group based on synapomorphies of the wings, abdominal sterna, female genitalia and the abdominal trichobothria.
The position of the Aradoidea is more contentious. Leston et al. (1954) indicated a possible relationship between aradoids and the 'Trichophora'. Schuh (1979, 1986) and Wheeler et al. (1993) provided morphological and molecular information to support this arrangement. Henry (1997) also argued for the inclusion of the aradoids in the Pentatomomorpha by synapomorphies of the pronotum and female ovipositor. Sweet (1996), however, postulated that the Aradoidea are not the sister-group of the trichophoran Pentatomomorpha, and warrant infraordinal ranking; he erected a new infraorder, the Aradomorpha, for their inclusion. Schaefer (1993) also alluded to possible alternative arrangements for the Aradoidea. Neither worker provided a strong evidential basis for their proposals. Schuh (1996) rejected these arguments for an alternative arrangement of the Aradoidea.
Schuh (1986) and Schaefer (1993) summarised the history of the classification of the Pentatomomorpha.
Superfamily Classification
The monophyly, sister-group relationships and composition of the pentatomomorphan superfamilies have been the subject of alternative views (Cassis & Gross 2002: Table 1). Six or seven superfamilies are recognised in the literature.
In summary, the alternative classifications vary in the superfamilial position of a number of families (Idiostolidae, Piesmatidae and Thaumastellidae), the validity of the Pyrrhocoroidea, and the concept of the Lygaeoidea and Coreoidea. Štys & Kerzhner (1975) held a broad view of the Coreoidea which included the Lygaeoidea of others. However, nearly all subsequent authors have regarded these two latter taxa as distinct.
Central to this problem has been the poor definition of many of the family groups, some of which are possibly paraphyletic. Henry (1997) established the monophyly of the families of the Lygaeoidea, which has resulted in considerable subdivision of the Lygaeidae and the recognition of 10 families not previously accepted by previous modern workers (cf. classifications in Cassis & Gross 2002: Table 1). His work is a radical departure from all other alternatives but has very strong evidential support and is gaining acceptance by key workers in the field (e.g. Slater 1998). Henry (1997) also provided phylogenetic support for the other superfamilies and the included families. His classification was followed in Cassis & Gross (2002).
TAXONOMIC WORKS
Catalogues, Checklists and Bibliographies
Stål (1870–1876) and Lethierry & Severin (1893–1896) provided the only catalogues of the Pentatomomorpha of the world (the latter updated by Bergroth 1908, 1913). Kirkaldy (1909) catalogued the majority of the Pentatomoidea (Aphylidae, Dinidoridae, Pentatomidae, Phloeidae, Plataspidae, Scutelleridae and Tessaratomidae) and this work has been important in outlining the complex nomenclature of the Pentatomidae.
Modern bibliographic works are either at the family level or cover a particular region. Stonedahl & Dolling (1991) listed catalogues and checklists of the Pentatomomorpha. Schuh & Slater (1995) provided updated information on critical literature. The family-group catalogues and checklists that have been used are those of the Aphylidae (Schouteden 1906a), Aradidae (Usinger & Matsuda 1959; Kormilev & Froeschner 1987), Berytidae (Henry & Froeschner 1998), Cydnidae (Lis 1996), Dinidoridae (Schouteden 1913; Rolston et al. 1996), Largidae (Hussey 1929), Lygaeidae sensu lato (Slater 1964; Slater & O'Donnell 1995—also including Artheneidae, Blissidae, Cryptorhamphidae, Cymidae, Geocoridae, Heterogastridae, Ninidae, Oxycarenidae, Pachygronthidae and Rhyparochromidae), Pentatomidae (Schouteden 1905, 1907), Pyrrhocoridae (Hussey 1929), Rhopalidae (Göllner-Scheiding 1983), Scutelleridae (Schouteden 1904, 1906b) and Tessaratomidae (Rolston et al. 1993).
Dr David Rider (in litt.) provided us with critical support in developing the Pentatomidae section of Cassis & Gross (2002). He is presently preparing a catalogue of the Pentatomidae of the world that includes a new subfamilial and tribal classification. He gave us access to his manuscript, which we used extensively in the preparation of the 2002 Catalogue. This resulted in some new arrangements for genera in the tribes of the Pentatominae.
Regional catalogues and checklists were also been useful in the compilation of Cassis & Gross (2002). The majority of such catalogues of the Pentatomomorpha exist for the Western Hemisphere. The North American fauna has been catalogued by Van Duzee (1916, 1917) and Henry & Froeschner (1988, and cohorts). Froeschner (1981—Ecuador, 1985—Galápagos, 2000—Panama) provided partial checklists of the Heteroptera of the northern Neotropical region. Few catalogues and checklists of the Eastern Hemisphere are available that are relevant to the Catalogue. The Palaearctic fauna was treated by Oshanin (1906–1912) and Stichel (1955–1959), but these works have not been of great significance. Aukema & Rieger (1995, 1996) and others, for example Kerzhner & Josifov (1999) are cataloguing the entire Palaearctic fauna. These works were useful in verifying bibliographic material. Atkinson (1890) provided a catalogue of the Indian fauna, but this work has limited value. There are no modern catalogues and checklists of the Oriental and Afrotropical Regions. However, faunistic studies (e.g. Linnavuori 1978) were useful in understanding the Pentatomomorpha of these regions.
As in Cassis & Gross (1995), we used the bibliography of Musgrave (1930) extensively and also his unpublished materials held by the Australian Museum.
Taxonomic Revisions and Activity
The numbers of Pentatomomorpha genera and species for Australia as at 2002 were given in Cassis & Gross (2002: Table 2). An indication of the taxonomic effort over particular periods can be deduced from the graph of cumulative number of heteropteran species given for Australia by Cassis & Gross (1995). The most significant activity has occurred over the past fifty years. In the first 100 years post-Linnaeus, activity was limited, a mere 200 species being described. In the next 50 years there was considerable work on the Australian fauna by workers at the British Museum of Natural History (now The Natural History Museum, London), with Dallas, Walker and Distant describing many new species.
Up to the middle of the last century about 650 species of Pentatomomorpha had been described, a mere 200 more than at the beginning of that century. This was a period of limited taxonomic activity on the Australian Pentatomomorpha. For the Australian fauna, the most active period has been the last fifty years, over which 500 new species have been described. The most significant contributions have been to the Aradoidea (Kormilev, Monteith), Coreoidea (Brailovsky), Lygaeoidea (Gross, Malipatil, Scudder, Slater, Woodward) and Pentatomoidea (Gross, McDonald).
REVIEW OF THE AUSTRALIAN PENTATOMOMORPHAN FAUNA
Composition of the Australian Pentatomomorphan Fauna
The described Australian Pentatomomorpha now comprises 1203 species, in 34 families and 449 genera. The fauna has a notable autochthonous component with four endemic families: Henicocoridae, Hyocephalidae, Aphylidae and Lestoniidae. A number of tribes are either endemic (Coreidae: Agriopocorini) or found chiefly in Australia (Coreidae: Amorbini and Rhyparochromidae: Udeocorini).
The fauna is strongly endemic at both the generic and species levels, with 52% of the genera and 83% of the species restricted to Australia. This degree of insularity of the Pentatomomorpha is remarkably similar to the lower Heteroptera (Coleorrhyncha-Cimicomorpha), at 45% of genera and 83% of species (Cassis & Gross 1995). The Pentatomidae have a very high level of generic endemicity, at over 70%. In many of the larger families (Aradidae, Lygaeidae, Rhyparochromidae, Coreidae, Acanthosomatidae, Cydnidae and Tessaratomidae) half (or thereabouts) of the genera are endemic to Australia. Many of the smaller families that are found extralimitally are represented in Australia by endemic genera alone; these include the Idiostolidae, Artheneidae, Cryptorhamphidae and Piesmatidae.
Species endemicity, as in the Coleorrhyncha-Cimicomorpha, is very high for the majority of pentatomomorphan families. Of the speciose families (Aradidae, Lygaeidae, Rhyparochromidae, Coreidae, Acanthosomatidae, Cydnidae and Pentatomidae), at least 75% of the species are restricted to Australia. For a number of the smaller families species endemicity is also high, and this is true for the Termitaphididae, Artheneidae, Berytidae, Blissidae, Colobathristidae, Cryptorhamphidae, Geocoridae, Heterogastridae, Piesmatidae, Rhopalidae and Pyrrhocoridae.
Monteith (1997) suggested that the Australian Aradidae: Mezirinae fauna is nearing completion in terms of discovery and description. This is also likely to be the situation for some of the smaller families such as the Termitaphididae, Idiostolidae, Henicocoridae, Colobathristidae, Cryptorhamphidae, Cymidae, Geocoridae, Heterogastridae, Ninidae, Oxycarenidae, Pachygronthidae, Largidae, Pyrrhocoridae, the coreoid families, Aphylidae, Dinidoridae, Lestoniidae, Plataspidae, Scutelleridae and Tessaratomidae. Although there has been considerable taxonomic work on the two most diverse pentatomomorphan families, the Rhyparochromidae and Pentatomidae, further collecting in more xeric habitats will undoubtedly result in the discovery of new taxa.
Schuh & Slater (1995) and others (see family introductions) have estimated components of the Heteropteran fauna on a worldwide basis. We estimate the world Pentatomomorpha fauna to comprise 2347 genera and 15,405 species. On these figures the Australian pentatomomorphan fauna represents 18% of the genera and 8% of the species of the world. These figures are commensurate with the notion that Australia is one of the megadiverse nations of the world. This is further supported by the fact that the study of the Australian Heteroptera is still within an age of discovery and description, and in all likelihood the fauna is considerably more diverse.
Distribution
The Pentatomomorpha are found in all States and territories of Australia (Table 3). As in the Cimicomorpha-Coleorrhyncha, the majority of pentatomomorphan species are found on the eastern seaboard. The infraorder is most diverse in Queensland, particularly in the wet tropics, and currently two-thirds (779 species) of the Australian Pentatomomorpha are found in this State. The New South Wales fauna comprises 397 species, about a third of the Australian fauna, with the majority of species found in the coastal regions. Species richness drops significantly in the cool temperate regions of eastern Australia, particularly in Victoria (253 species) and Tasmania (126 species). The Pentatomomorpha of South Australia (325 species) is almost as diverse as the New South Wales fauna, but this is partly confounded by the extensive collecting and taxonomic work on the State's fauna by Gross (e.g. 1975, 1976). The described Western Australian pentatomomorphan fauna is relatively depauperate (288 species), particularly in relation to the geographic size of that State. However, this is undoubtedly an under-representation. One of us [GC] has made significant collections in the south-western floristic region of Western Australia, an area noted for its biotic endemism and diversity, and these have revealed a very species rich and understudied heteropteran fauna.
The biogeographic relationships of the Australian pentatomomorphan fauna are poorly understood. Trans-oceanic distribution patterns of these bugs can be described as either Indo-Pacific (e.g. Catacanthus Spinola), Indo-Malayan (e.g. Cuspicona Dallas), Australian (east of Wallace Line, e.g. Chiastoplonia China) or Gondwanan (e.g. Idiostolidae). The most significant feature of the biogeography of the Australian Pentatomomorpha is its highly insular nature, which is reflected in the high percentage of endemic taxa (Cassis & Gross 2002: Table 2). Most attempts to define the area relationships are based on narratives of the distribution patterns and no historical biogeographic analyses have been published.
The Pentatomomorpha are represented on all the Australian territorial islands, aside from the Subantarctic islands (Cassis & Gross 2002: Tables 3 and 4). The Christmas Island fauna contains 22 species, 9 of which are endemic, and are best represented by the Rhyparochromidae. The Lord Howe Island fauna comprises fifteen described species (4 endemic species), but is known to be far more diverse, with many new endemic species awaiting description. Norfolk Island and Cocos-Keeling Islands have eight and four species respectively.
Host Associations
The majority of Pentatomomorpha are phytophagous and are either mycetophagous (Aradoidea), granivorous (most of Lygaeoidea, Coreoidea, Pentatomoidea and Pyrrhocoroidea) or folivorous and/or fructivorous (most Pentatomoidea, but also feed on stems, cambium and petioles). Schaefer & Panizzi (2000) reviewed extensively the biology of many of the Pentatomomorpha, with particular reference to species of economic importance. Their work contains an exhaustive list of biological references, and we have avoided any duplication of their effort here. It is beyond the scope of this Catalogue to review adequately the biology of the Australian Pentatomomorpha or the species of economic significance.
The discussion that follows is restricted intentionally to a narrative on host associations. Important reference materials that pertain to host associations, in addition to Schaefer & Panizzi (2000), include: Ahmad & Schaefer (1987—Pyrrhocoroidea), Schaefer & O'Shea (1979—Coreidae), Schaefer (1980—Alydidae), Schaefer & Chopra (1982—Rhopalidae), Schaefer (1983—Piesmatidae and Podopinae), Schaefer & Mitchell (1983—Coreoidea), Schaefer & Ahmad (1987—Acanthosomatidae, Tessaratomidae, Urostylidae and Dinidoridae) and Schaefer (1988—Pentatomoidea).
The host associations of phytophagous Pentatomomorpha are given in Tables 5, 6, 7 and Appendix IV. There are 855 host plant records for the Australian Pentatomomorpha. The recorded host plant species belong to 29 plant orders and 82 plant families. The following discussion is based on the plant phylogeny and classification of The Angiosperm Phylogeny Group (1998). Note that in the following discussion reference to rosids and asterids refers to rosids + eurosids and asterids + euasterids respectively.
The pentatomomorphan host records suggest a strong pattern of association between the Pentatomomorpha and the asterid and rosid families of eudicotylodenous angiosperms. The infraorder appears not to exploit the gymnosperms of Australia to any great extent (Table 5), with only four records (Rhyparochromidae and Pentatomidae) in the Catalogue. The pentatomomorphans are essentially not represented on the basal angiosperms (Laurales and Magnoliales), with the only records existing for Amblypelta Stål species (Coreidae) feeding on commercial fruits; avocados (Lauraceae) and custard apples (Annonaceae) (Table 5).
The monocots are exploited by 15 families of Pentatomomorpha (in Coreoidea, Lygaeoidea and Pentatomoidea) (Table 5). On the basis of current information, these true bugs use plants belonging to the Poales more than any other plant order (66 records), aside from the Myrtales. A number of pentatomomorphans are grass-specialists—Blissidae, Colobathristidae and Pachygronthidae. The Cryptorhamphidae, Cymidae and Ninidae feed on a broader range of Poales, including grasses, sedges, rushes and restios. Grass-feeding also occurs in coreoids, with the Alydidae: Leptocorisina being most notable. In the Pentatomidae there are several grass associations and it is possible that the Pentatominae: Aeptini are specialist grass-feeders.
The basal eudicots (Core Eudicots, Ranunculales, Proteales, Caryophyllales, Santalales and Saxifragales) do not appear to be utilised by the Pentatomomorpha to any large extent (Table 6). Coreidae are known from seven families of basal eudicots with most host records on Proteaceae, Nyctaginaceae, Polygonaceae and Grossulariaceae. Pentatomidae are also known from a range of basal eudicot families, and particularly from Proteaceae and Chenopodiaceae.
The range and frequency of association of pentatomomorphan families is highest with the rosid angiosperms. Nineteen bug families are recorded from 34 families and over 500 plant records (Table 7). Pentatomomorphan species are known from all the rosid orders, with most records from species in the Fabales and Myrtales. Lygaeidae, Rhyparochromidae, Coreidae and Pentatomidae are well represented on Fabaceae, Mimosaceae, Cucurbitaceae, Myrtaceae and Rutaceae. The Myrtaceae are the plant family most exploited by the Pentatomomorpha, with 173 records for the infraorder. The plant genera that are most utilised are Eucalyptus L'Hér (42 species), Leptospermum J.R.&G. Forst. (4 species) and Melaleuca Linnaeus (10 species). In general, not many suprageneric groups of Heteroptera have exploited Eucalyptus to any great extent, unlike the Auchenorrhyncha. However, in the Pentatomomorpha, the coreid genus Amorbus Dallas and several species of the Pentatominae: Halyini (particularly Poecilometis Dallas) are clearly adapted to life on gum trees. The Pentatomomorpha are also often associated with species of Acacia Mill. (Mimosaceae), sennas (Cassia Linnaeus) and a variety of legumes (Fabaceae). Pentatomomorphans are recorded from 22 species of Acacia, but we consider this to be a gross under-sampling, and wattles may turn out to be the plant group most heavily utilised by this group of bugs in Australia.
The Pentatomomorpha of Australia have not been as successful at utilising the asterids as they have the rosids. Twelve families of pentatomomorphans are recorded from 19 asterid eudicot families (with 168 host plant records) (Table 8). Lygaeidae (particularly Lygaeinae) have exploited asterids more than any other pentatomomorphan family, and are mostly found on Apocynaceae, Asclepiadaceae, Solanaceae and Asteraceae. Coreidae are known from ten asterid families (18 host species records), with the most frequent association occurring with Solanaceae. There are 48 host records (10 families) for the Pentatomidae, and they also are most often associated with Solanaceae, and then most frequently with Asteraceae and Myoporaceae.
A number of families exhibit very broad host ranges, particularly the Lygaeidae (15 plant orders), Rhyparochromidae (17), Coreidae (24) and Pentatomidae (22). These results are somewhat confounded by polyphagous species, such as Nysius vinitor Bergroth, Leptoglossus gonagra (Fabricius) and Nezara viridula (Linnaeus), which are known from many unrelated plants. However, the degree of host specificity for most pentatomomorphan species is high, as half of the recorded associations are with a single plant species.
The animal prey of predacious pentatomomorphans are very poorly known. The majority of records are for the asopine species Andrallus spinidens (Fabricius), Cermatulus nasalis (Westwood) and Oechalia schellenbergi (Guérin) which feed mostly on the larvae of Lepidoptera.
Acknowledgements
Many Heteropterists have greatly assisted in the preparation of this work. Special thanks are given to Dr David Rider who freely provided unpublished material on the Pentatomidae and very detailed reviews of much of the Catalogue. We greatly appreciate his generosity and acknowledge the originality of his ideas. This Catalogue would have been greatly compromised without his assistance.
Dr Geoff Monteith and Mr Tom Weir provided significant support throughout the duration of this work, both in terms of information, review of the manuscript, and encouragement. Their support is warmly appreciated. Mr Lionel Hill responded without restraint to calls for assistance. Drs Martin Steinbauer and Mali Malipatil reviewed the Coreoidea and Lygaeoidea sections of the Catalogue and are duly thanked. Dr Jerzy Lis provided us with critical information at the 11th hour; his help is much appreciated.
Dr Toby Schuh was of invaluable assistance in providing literature, reviewing the manuscript, and giving access to his vast knowledge of the world Heteroptera. Dr Jim Slater gave us similar support for the Lygaeoidea. Drs Dick Froeschner and Carl Schaefer were very generous in providing reprints of their work.
This project was also greatly assisted by museum curators across Australia. We wish to thank Mr Greg Daniels, Mr John Donaldson, Dr Terry Houston, Mr Geoff Thompson, Dr Murray Fletcher and Dr Ken Walker.
The preparation of this volume would not have been possible without the great help of staff at the Australian Museum Centre for Biodiversity and Conservation Research. Ms Rossana Silveira deserves special mention for her skill and diligence in seeing this project to its completion. She entered data, sourced bibliographic materials, prepared the graphs—all done with good humour and friendship. Mr Michael Elliot was also a great help with many aspects of the Catalogue and we acknowledge his assistance. Dr Lance Wilkie was very helpful in analysing the host information. Other staff members who helped in many ways include Dr Max Moulds, Mr Barry Day, Mr Gareth Carter, Dr David McAlpine and Ms Sue Lindsay. We would also like to make special mention of Dr Chris Reid who listened to endless discussion about 'the catalogue'. He never looked bored and we thank him for his advice.
CSIRO Entomology and Melbourne University Press are thanked for providing the majority of the habitus illustrations. Other people who have kindly provided habitus illustrations include Dr Mali Malipatil (Oxycarenidae) and Ms Jan Forrest (Aphylidae and Scutelleridae). Ms Hannah Findlay prepared fine new illustrations (Artheneidae, Cryptorhamphidae, Cymidae, Henicocoridae, Heterogastridae and Ninidae) under our supervision and her efforts are acknowledged.
We would like to thank the staff of the Australian Biological Resources Study for their help in bringing closure to this project. We especially thank Drs Keith Houston and Alice Wells for their editorial skills and time.
We thank our families for their encouragement, support and patience.
Database Notes
The Catalogue is arranged alphabetically by superfamily, with the families therein arranged alphabetically. All genera and species are arranged alphabetically within families. Taxa that have uncertain taxonomic position are either listed as Unplaced within a higher group, or as incertae sedis at the end of a family treatment.
The methods employed in the compilation of the Catalogue are outlined in Cassis & Gross (1995). Rule changes given in IZCN (1999) have been incorporated in the preparation of the Catalogue.
30 September 2002
Edited out extra extralimital distribution data from file, put in page numbers for 'this work' in hard copy, deleted 'new record/s' from distribution data. Ready for loading on Web.
Limital Area
Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.
Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.
Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124 DEG E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.
Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.
Excluded Taxa
- Misidentifications
ARADIDAE: Arictus chinai (Kormilev, 1955)
SCUTELLERIDAE: Calliphara nobilis (Linnaeus, 1763)
SCUTELLERIDAE: Calliphara regia (Westwood, 1837)
SCUTELLERIDAE: Cantao africanus Horváth, 1892
CYDNIDAE: Cydnus aterrimus Forster, 1771
PENTATOMIDAE: Hyrmine sexpunctatus (Linnaeus, 1758)
CYDNIDAE: Macroscytus subaeneus Dallas, 1851
CYDNIDAE: Macroscytus transversus Burmeister, 1834
LYGAEIDAE: Melanotelus bipunctata Dallas, 1852
TESSARATOMIDAE: Plisthenes merianae (Fabicius, 1775)
RHYPAROCHROMIDAE: Poeantius lineatus Stål, 1874
LYGAEIDAE: Spilostethus pandurus elegans Wolff, 1802
General References
Ahmad, I. & Schaefer, C.W. 1987. Food plants and feeding biology of the Pyrrhocoroidea (Hemiptera). Phytophaga. Palermo 1: 75-92
Atkinson, E.T. 1890. Catalogue of the Insecta. Order Rhynchota. Suborder Hemiptera-Heteroptera. Family Capsidae. Journal of the Asiatic Society of Bengal Nat. Sci. Suppl. 58: 25-200
Bergroth, E. 1908. Enumeratio Pentatomidarum post Catalogum bruxellensem descriptarum. Mémoires de la Société Entomologique de Belgique 15: 131-200
Bergroth, E. 1913. Supplementum Catalogi Heteropterorum Bruxellensis II. Coreidae, Pyrrhocoridae, Colobathristidae, Neididae. Mémoires de la Société Entomologique de Belgique 22: 125-183
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. 1995. Hemiptera: Heteroptera (Coleorrhuncha to Cimicomorpha). pp. 1-501 in Houston, W.W.K. & Maynard, G.V. (eds). Zoological Catalogue of Australia. Hemiptera: Coleorrhyncha to Cimicomorpha. Melbourne : CSIRO Australia Vol. 27.3A xv 506 pp.
Dolling, W.R. 1991. The Hemiptera. Oxford : Oxford University Press ix 274 pp.
Froeschner, R.C. 1981. Heteroptera or true bugs of Ecuador: a partial catalog. Smithsonian Contributions to Zoology 1981(322): iv 1-147
Froeschner, R.C. 1985. Synopsis of the Heteroptera or true bugs of the Galápagos Islands. Smithsonian Contributions to Zoology 407: iii 1-84
Froeschner, R.C. 2000. True Bugs (Heteroptera) of Panama: A synoptic catalog as a contribution to the study of Panamanian biodiversity. Memoirs of the American Entomological Institute 62: ii 1-393
Göllner-Scheiding, U. 1983. General-Katalog der Familie Rhopalidae (Heteroptera). Mitteilungen aus dem Zoologischen Museum in Berlin 59: 37-189
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
Henry, T.J. & Froeschner, R.C.. 1998. Catalog of the Stilt Bugs or Berytidae, of the World (Insecta: Hemiptera: Heteroptera). Contributions of the American Entomological Institute 30(4): 1-72
Kerzhner, I.M. & Josifov, M. 1999. Catalogue of Palaearctic Heteroptera. Cimicomorpha I. pp. xiv 1-577 in Aukema, B. & Rieger, C. (eds). Catalogue of the Heteroptera of the Palaearctic Region. Wageningen : Ponsen & Looijen Vol. 3 xiv 577 pp.
Kormilev, N.A. & Froeschner, R.C. 1987. Flat bugs of the world: a synonymic list (Heteroptera: Aradidae). Entomography 5: 1-246
Leston, D., Pendergrast, J.G. & Southwood, T.R.E. 1954. Classification of the terrestrial Heteroptera (Geocorisae). Nature (London) 174: 91-92
Linnavuori, R. 1978. Hemiptera of the Sudan, with remarks on some species of the adjacent countries 6. Aradidae, Meziridae, Aneuridae, Pyrrhocoridae, Stenocephalidae, Coreidae, Alydidae, Rhopalidae, Lygaeidae. Acta Zoologica Fennica 153: 1-108
Lis, J.A. 1996. A review of burrower bugs of the Australian Region, with a discussion on the distribution of the genera (Hemiptera: Heteroptera: Cydnidae). Genus (Wroclaw) 7(2): 177-238
Monteith, G.B. 1997. Revision of the Australian flat bugs of the subfamily Mezirinae (Insecta: Hemiptera: Aradidae). Memoirs of the Queensland Museum 41(1): 1-169
Rolston, L.H., Aalbu, R.L., Murray, M.J. & Rider, D.A. 1993. A catalog of the Tessaratomidae of the world. Papua New Guinea Journal of Agriculture, Forestry and Fisheries 36: 36-108
Rolston, L.H., Rider, D.A., Murray, M.J. & Aalbu, R.L. 1996. Catalog of the Dinidoridae of the World. Papua New Guinea Journal of Agriculture, Forestry and Fisheries 39(1): 22-101
Schaefer, C.W. 1975. Heteropteran trichobothria, (Hemiptera-Heteroptera). International Journal of Insect Morphology and Embryology 4: 193-264
Schaefer, C.W. 1980. The host plants of Alydinae, with a note on heterotypic feeding aggregations (Hemiptera: Coreoidea: Alydidae). Journal of the Kansas Entomological Society 53: 115-122
Schaefer, C.W. 1983. Host plants and morphology of the Piesmatidae and Podopinae (Hemiptera: Heteroptera): further notes. Annals of the Entomological Society of America 76: 134-137
Schaefer, C.W. 1988. The food plants of some "primitive" Pentatomoidea (Hemiptera: Heteroptera). Phytophaga. Palermo 2: 19-45
Schaefer, C.W. 1993. The Pentatomomorpha (Hemiptera: Heteroptera): an annotated outline of its systematic history. European Journal of Entomology 90: 105-122
Schaefer, C.W. & Ahmad, I. 1987. The food plants of four pentatomoid families (Hemiptera: Acanthosomatidae, Tessaratomidae, Urostylidae, and Dinidoridae). Phytophaga. Palermo 1: 21-34
Schaefer, C.W. & Chopra, N.P. 1982. Cladistic analysis of the Rhopalidae, with a list of food plants. Annals of the Entomological Society of America 75: 224-233
Schaefer, C.W. & Mitchell, P.L. 1983. Food plants of the Coreoidea (Hemiptera: Heteroptera). Annals of the Entomological Society of America 76: 591-615
Schaefer, C.W. & O'Shea, R. 1979. Host plants of three coreine tribes (Hemiptera: Heteroptera: Coreidae). Annals of the Entomological Society of America 72: 519-523
Schouteden, H. 1904. Heteroptera. Fam. Pentatomidae. Subfam. Scutellerinae. Genera Insectorum 24: 1-98 5 pls
Schouteden, H. 1905. Heteroptera. Fam. Pentatomidae. Subfam. Graphosomatinae. Genera Insectorum 30: 1-46 3 pls
Schouteden, H. 1906a. Heteroptera. Fam. Pentatomidae. Subfam. Aphylinae. Genera Insectorum 47: 1-4 1 pl.
Schouteden, H. 1906b. Heteroptera. Fam. Pentatomidae. Subfam. Scutellerinae. Genera Insectorum 24: 99-100
Schouteden, H. 1907. Heteroptera. Fam. Pentatomidae. Subfam. Asopinae (Amyoteinae). Genera Insectorum 52: 1-82 5 pls
Schouteden, H. 1913. Heteroptera. Fam. Pentatomidae. Subfam. Dinidorinae. Genera Insectorum 153: 1-19 2 pls
Schuh, R.T. 1979. Book review of Evolutionary Trends in Heteroptera. Part II. Mouthpart-structures and Feeding Strategies.—R.H. Cobben. 1978. Meded. Landbouwhogeschool Wageningen 78–5. 407 pp. Systematic Zoology 28: 653-656
Schuh, R.T. 1986. The influence of cladistics on Heteropteran classification. Annual Review of Entomology 31: 67-93
Schuh, R.T. 1996. Book Review of "C.W. Schaefer (ed.) 1996. Studies on Hemipteran Phylogeny. Proceedings. Thomas Say Publications on Entomology. 244 pp. Entomological Society of America. Lanham, Maryland". Journal of the New York Entomological Society 104(3–4): 231-235
Slater, J.A. 1998. New genera and species of Lygaeoidea (Heteroptera: Lygaeoidea: Rhyparochromidae). Insecta Mundi 12(3–4): 239-250
Stål, C. 1870. Enumeratio Hemipterorum. Bidrag till en förteckning öfver aller hittills kända Hemiptera, jemte systematiska meddelanden. 1. Kongliga Svenska Vetenskaps-Academiens Nya Handlingar, Stockholm n.f. 9(1): 1-232
Stål, C. 1872. Enumeratio Hemipterorum. Bidrag till en förteckning öfver aller hittills kända Hemiptera, jemte systematiska meddelanden. 2. Kongliga Svenska Vetenskaps-Academiens Nya Handlingar, Stockholm n.f. 10(4): 1-159
Stål, C. 1873. Enumeratio Hemipterorum. Bidrag till en förteckning öfver aller hittills kända Hemiptera, jemte systematiska meddelanden. 3. Kongliga Svenska Vetenskaps-Academiens Nya Handlingar, Stockholm n.f. 11(2): 1-163
Stål, C. 1874. Enumeratio Hemipterorum. Bidrag till en förteckning öfver aller hittills kända Hemiptera, jemte systematiska meddelanden. 4. Kongliga Svenska Vetenskaps-Academiens Nya Handlingar, Stockholm n.f. 12(1): 1-186
Stål, C. 1876. Enumeratio Hemipterorum. Bidrag till en förteckning öfver aller hittills kända Hemiptera, jemte systematiska meddelanden. 5. Kongliga Svenska Vetenskaps-Academiens Nya Handlingar, Stockholm n.f. 14(4): 1-162
Stichel, W. 1955. Illustrierte Bestimmungstabellen der Deutschen Wanzen. II. Europa. (Hemiptera-Heteroptera Europae). Berlin-Hermsdorf : Selbstverlag pp. 1-4. [Vol. 1: 1–168; Vol. 2: 170–907; Vol. 3: 1–428; Vol. 4: 1–838]
Stichel, W. 1958. Nomenklatorisches über Heteropteren. Bollettino della Società Entomologica Italiana 88: 21-23
Stichel, W. 1959. Illustrierte Bestimmungstabellen der Deutschen Wanzen. II. Europa (Hemiptera-Heteroptera Europae). Berlin-Hermsdorf : Verlag W. Stichel Vol. 4(8-12) pp. 225-384. [Pt 8, Pentatomomorpha Lygaeoidea Lygaeidae (7) pp. 225–256; Pt 9, Pentatomomorpha Lygaeiodea Lygaeidae (8) pp. 257–288; Pt 10, Pentatomomorpha Lygaeidae (9), Pyrrhocoridae Addenda, Corrigenda Supplementa Liste der paläarktischen Arten (1) pp. 289–320; Pt 11, Liste der paläarkstischen Arten (2. Fortsetzung) Pentatomomorpha Lygaeidae pp. 321–352; Pt 12, Pentatomomorpha Coreoidea Stenocephalidae Coreidae (Coreinae) pp. 353–384]
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. & Kerzhner, I. 1975. The rank and nomenclature of higher taxa in recent Heteroptera. Acta Entomologica Bohemoslovaca 72: 65-79
Sweet, M.H. 1996. Comparative External Morphology of the Pregenital Abdomen of the Hemiptera. pp. 119-157 in Schaefer, C.W. (ed.). Studies on Hemipteran Phylogeny. Lanham : Entomological Society of America 244 pp.
The Angiosperm Phylogeny Group 1998. An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Gardens 85: 531-553
Tullgren, A. 1918. Zur Morphologie und Systematik der Hemipteren I. Entomologisk Tidskrift 1918: 113-133
Van Duzee, E.P. 1917. Catalogue of the Hemiptera of America north of Mexico, excepting the Aphididae, Coccidae and Aleurodidae. University of California Publications in Entomology. Technical Bulletin 2: xiv 1-902
Wheeler, W.C., Schuh, R.T. & Ranhy Bang 1993. Cladistic relationships among higher groups of Heteroptera: congruence between morphological and molecular data sets. Entomologica Scandinavica 24: 121-137
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.
History of changes
Published | As part of group | Action Date | Action Type | Compiler(s) |
---|---|---|---|---|
05-Dec-2019 | PENTATOMOMORPHA | 05-Dec-2019 | MODIFIED | |
15-Aug-2012 | 04-Oct-2019 | MODIFIED | ||
05-Dec-2019 | MODIFIED |