Gerrids, Reduviids, Water-striders
Compiler and date details
June 2012 - Professor Gerry Cassis, Anna Namyatova, Nikolai Tatarnic and Celia Symonds, University of New South Wales, Sydney
30 June 1995 - Gerasimos Cassis, Australian Museum, Sydney, Australia; Gordon Gross, South Australian Museum, Adelaide, Australia
Note: The Heteroptera (Coleorrhyncha to Cimicomorpha) section of the Australian Fauna Directory, is derived from (Cassis, G. & Gross, G.F., 1995). It was transferred from the Zoological Catalogue of Australia database to the ABRS Platypus software program and prepared for the Australian Fauna Directory by Kathy Tsang and Keith Houston. It is currently being updated but is included in the Directory now in response to requests.
Users should note, however, that parts, e.g. Distribution and maps, have not been fully converted to the Platypus/AFD format pending the update, and tables and figures, cited in the Introductions and in the original published work, are not included.
Hemiptera are the largest and most diverse of the exopterygote orders of insects. They are suction feeders and include groups commonly known as true bugs, aphids, psyllids, whiteflies, scale insects, leafhoppers, planthoppers, froghoppers and cicadas.
The order is grouped in the subdivision Paraneoptera (= Acercaria, hemipteroid assemblage), with the orders Psocoptera, Phthiraptera and Thysanoptera (Hennig 1981; Kristensen 1991; Kukalová-Peck 1991). It consists of three suborders: Sternorrhyncha, Auchenorrhyncha and Heteroptera (Carver et al. 1991). Of these, the Heteroptera are the most diverse taxonomically and include aquatic, semiaquatic and terrestrial representatives. Depending on the classification (cf. Schuh 1986; Carver et al. 1991; Stonedahl & Dolling 1991; Table 1), this suborder comprises between 73 and 76 families, and over 39,000 species worldwide (Slater 1982; Zoological Record 1980-1994).
The Heteroptera section of the Zoological Catalogue of Australia is divided into two parts. Volume 27.3A (this work) covers the infraorders Coleorrhyncha, Enicocephalomorpha, Dipsocoromorpha, Gerromorpha, Leptopodomorpha, Nepomorpha and Cimicomorpha (hereafter referred to as Coleorrhyncha-Cimicomorpha). Volume 27.3B (Cassis & Gross, in preparation) covers all taxa in the Pentatomomorpha. Only extant taxa are included in the Catalogue, among them the Peloridiidae, which has often been placed outside the suborder. The Australian heteropteran fauna presently totals 55 families, 779 genera and 2006 species, of which 32 families, 352 genera and 909 species are covered in this section (Vol. 27.3A) of the Catalogue (see Table 1).
In the Catalogue, we have included all extralimital generic and species synonymies, as currently accepted in the literature. Where alternative taxonomic arrangements exist, the divergent views are listed. New synonymies, combinations, Australian records and lectotype designations are included. All relevant literature from 1758 to 30 June 1995 has been included in this volume. The family introductions include a review of the world fauna, morphology, biology and systematics, as well as a synopsis of the Australian fauna. For a general account of the morphology and biology of the Heteroptera, users are referred to Carver et al. (1991) and Dolling (1991).
The suprageneric classification utilised in the Catalogue basically conforms with that outlined in The Insects of Australia (Carver et al. 1991). It differs only in the superfamilial arrangement of Cimicomorpha, for which we follow the cladistic classification of Schuh & Stys (1991), which has more heuristic definitions of the Cimicoidea and Miroidea. The infraordinal, superfamilial and familial classifications used for both Heteroptera volumes of the Catalogue are outlined in Tables 1 and 2. The subfamilial and tribal classifications used in this work are listed in the Table of Contents.
Subordinal and Infraordinal Classifications
The Hemiptera are considered here to comprise three suborders: the Sternorrhyncha (aphids, psyllids, whiteflies, scale insects), Auchenorrhyncha (leafhoppers, planthoppers, froghoppers, cicadas) and Heteroptera (true bugs). This classification is a recent departure from the traditional split of Homoptera (= Coleorrhyncha + Sternorrhyncha + Auchenorrhyncha) and Heteroptera but has received considerable support (Carver et al. 1991; Kukalová-Peck 1991; Dolling 1991, in part; Wheeler et al. 1993). These authors concluded that the Homoptera are not monophyletic and consequently raised the Sternorrhyncha and Auchenorrhycha to subordinal level, recognising that the latter are more closely related to the Heteroptera.
The position of the Peloridiidae within the Hemiptera is contentious. China (1924) suggested the erection of a separate suborder, the Pseudohomoptera, to accommodate this family. Myers & China (1929) placed them in the Homoptera on the basis of plesiomorphic characters. This arrangement was accepted by most authors until China (1962), in a thorough review of peloridiid morphology, suggested they be placed within the Auchenorrhyncha in their own infraorder, the Peloridiidomorpha (= Coleorrhyncha). Schlee (1969) and Hamilton (1981) recognised a sister-group relationship between Heteroptera and peloridiids. Emeljanov (1987) supported this arrangement, based on wing characters, although he disputed the presence of a gula in peloridiids. Carver et al. (1991) recently reassessed the affinities of Peloridiidae and placed them in the Heteroptera on the basis of wing structure and venation, presence of a gula, keeled propleura, trichobothria, cryptocerate antennae and mycoid microsculpture. Schlee (1969) proposed an infraordinal category, the Heteropteroidea (emended to Heteropterodea by Zrzavy 1990), comprising the Coleorrhyncha and Heteroptera. This classification is supported by Wheeler et al. (1993), who refrain from placing the Peloridiidae within the Heteroptera, presumably because of complications in re-defining the Heteroptera phylogenetically. In the Catalogue, we refer the Peloridiidae to their own infraordinal group (Coleorrhyncha) within the Heteroptera. This is to maintain consistency with the classification used in The Insects of Australia, despite the polemics that remain, and we refrain from supporting the use of the Heteropterodea nomenclature.
Traditionally, the Heteroptera were divided into three major groups, the Geocorisae (terrestrial bugs), the Hydrocorisae (aquatic bugs, also called Cryptocerata) and the Amphibiocorisae (water-surface dwelling bugs). The higher classification of the Heteroptera has recently received considerable attention, much of it stemming from the influential work of Leston et al. (1954). They introduced the -morpha infraordinal groups, which were based more on phylogenetic lines, and further subdivided the terrestrial bugs (Cimicomorpha and Pentatomomorpha). Stys & Kerzhner (1975) provided a comprehensive classification of the suborder and recognised seven infraorders, the Enicocephalomorpha, Dipsocoromorpha, Gerromorpha, Leptopodomorpha, Nepomorpha, Cimicomorpha and Pentatomorpha. These infraordinal divisions have remained stable and have been supported by cladistic analyses of morphological (Schuh 1979) and molecular information (Wheeler et al. 1993). Alternatively, Mahner (1993), based on new evidence, supports a more traditional classification, viz. Cryptocerata (= Nepomorpha or Hydrocorisae), and Gymnocerata which he divides into Amphibiocorisae (= Gerromorpha) and Geocorisae for the terrestrial groups, except for the Enicocephalomorpha which he leaves as incertae sedis. Schuh (1986) gave a thorough review of the infraordinal categories and listed the constituent families. For infraordinal names not mentioned in the Catalogue, users are referred to Stys & Kerzhner (1975) who give a detailed synonymical list of higher taxonomic names.
Stys (1985) introduced additional categories for sister-group arrangements of infraordinal -morpha groups, as suggested, but not named, in a cladogram provided by Schuh (1979). These included the Euheteroptera (all Heteroptera excluding Enicocephalomorpha), the Neoheteroptera (also excluding the Dipsocoromorpha), and the Panheteroptera (also excluding the Gerromorpha). Wheeler et al. (1993) supported this classification and suggested other monophyletic groups for the Leptopodomorpha, Cimicomorpha and Pentatomomorpha, which they refrained from naming. In the Catalogue, we make no organisational use of these categories.
Stys & Kerzhner (1975) provided the most comprehensive review of the superfamilial classification of the Heteroptera. They used superfamilies extensively to group the families within a phylogenetic context below the infraordinal level. Schuh (1986) did not use superfamilies for the Enicocephalomorpha and Dipsocoromorpha and listed numerous cimicomorphan families as incertae sedis. We have maintained the superfamilial classification of The Insects of Australia (Carver et al. 1991), except for the Cimicomorpha. We have adopted the classification of Schuh & Stys (1991) who restricted the Cimicoidea to the Anthocoridae sensu lato, Plokiophilidae, Cimicidae and Polyctenidae. Gross & Cassis (in Carver et al. 1991) indicated the limits of the Cimicoidea as conjectural and tentatively placed the Nabidae in this group on the basis of extragenital insemination. Schuh & Stys (1991) placed the Nabidae and the extralimital Medocostidae in a new superfamily, the Naboidea, regarding the nabid type of insemination as independently derived. They placed the extralimital Velocipedidae, Microphysidae and Joppeicidae in their own superfamilies. The Miroidea was redefined to include the Thaumastocoridae, Miridae and Tingidae, and the Reduvioidea was retained as previously constituted (Reduviidae + Pachynomidae). Readers are referred to Slater (1982) for detailed descriptions of the superfamilies.
The number of heteropteran families recognised has increased significantly in the past thirty years since China & Miller (1959) recognised 52 families in their checklist of family groups. Stys & Kerzhner (1975) listed 73 families, a number which has more or less been accepted by subsequent authors (cf. Schuh 1986-76 families; Stonedahl & Dolling 1991-73 families; Dolling 1991-76 families; Carver et al. 1991-73 families). Certain groups of the Enicocephalidae (Stys 1989), Anthocoridae (Schuh & Stys 1991), Naucoridae (Stys & Jansson 1988) and the malcid line of lygaeoids (Schuh 1986) are ranked either as families or subfamilies. Despite accepting the superfamily re-arrangements of the Cimicomorpha proposed by Schuh & Stys (1991), we believe their division of the Anthocoridae, raising the Lasiochilinae and Lyctocorinae to family rank, is premature. Stys & Jansson (1988), in a review of the higher classification of the Nepomorpha, support the family ranking of the Aphelocheiridae and extralimital Potamocoridae, which are often considered as subfamilies of Naucoridae. We follow The Insects of Australia classification and the views of Polhemus (1979) and Polhemus & Polhemus (1988) and retain these groups as naucorid subfamilies. The family group classification of the Pentatomomorpha will be discussed in Volume 27.3B (Cassis & Gross, in preparation).
Catalogues, Checklists and Bibliographies
The only attempt to catalogue the world heteropteran fauna (excluding Nepomorpha and Miridae) was made by Lethierry & Severin (1893-1896). Their work was emended and updated by Bergroth (1908, 1913). No modern equivalent of this work exists. Stonedahl & Dolling (1991) give a detailed list of the many regional catalogues available, including those of restricted taxonomic coverage. The North American fauna has been catalogued recently by Henry & Froeschner (1988), who updated Van Duzee's (1917) total of 1,625 species to 3,834 species. Oshanin (1906-1912) catalogued the Palaearctic heteropteran fauna, as did Stichel (1955-1962), who also provided keys to European taxa. A five volume Catalogue of Palaearctic Heteroptera, by B. Aukema and C Rieger, is in preparation. No comprehensive catalogues cover any of the other major zoogeographical regions. Catalogues and checklists more restricted in their distributional range are available. Froeschner (1981, 1985) provided catalogues of the Ecuadorian islands including the Galápagos, which have some relevance for the remainder of the Neotropical heteropteran fauna. Only descriptive taxonomic works on the Oriental fauna exist. Distant's (1902-1918) descriptive works of the fauna of the Indian subcontinent remain the most significant and have some relevance for the Australian heteropteran fauna.
This section of the Catalogue is the first comprehensive list of the Heteroptera fauna of Australia. It is derived chiefly from the primary literature and world catalogues of restricted taxonomic coverage. These catalogues are those of the Miridae (Carvalho 1957-1960), Reduviidae (Maldonado Capriles 1990), Leptopodomorpha (Schuh et al. 1986), Aradidae (Kormilev & Froeschner 1987), Lygaeidae (Slater 1964), Pentatomoidea (Kirkaldy 1909) and Tingidae (Drake & Ruhoff 1960, 1965). The most useful checklists include Lundblad's work on the aquatic and semiaquatic Heteroptera of the Indo-Australian region (Lundblad 1933), an unpublished checklist of the Cimicoidea (Ford 1979), and checklists of the Cimicidae (Usinger 1966), Enicocephalidae (Stys 1977-including the Aenictopecheidae), Gelastocoridae (Todd 1961), Mesoveliidae (Horváth 1929), Naucoridae (La Rivers 1974), Nepomorpha (Stys & Jansson 1988), Peloridiidae (Evans 1981), Polyctenidae (Ferris & Usinger 1939), Rhopalidae (Göllner-Scheiding 1983) and Schizopteridae (Emsley 1969). Andersen's (1982) monograph of the world Gerromorpha is indispensible for students of semiaquatic bugs. It contains a complete list of family-group and genus-group names, and the most important literature.
Musgrave's (1930) bibliographic work on Australian insects was particularly useful in the compilation of the Catalogue, although it was of limited value for taxa with extralimital distributions. An unpublished bibliography of Australian entomology from 1930 to the late 1950's, and a card catalogue of the Australian Heteroptera, also compiled by Musgrave, were both extremely useful.
Taxonomic Revisions and Activity
The cumulative number of species of Australian Heteroptera is outlined in Graph 1. Work on Heteroptera is a reflection of the history of Australian entomology in general and can be divided into separate periods of activity (Marks 1991). These are recognised as the Fabrician (1770-1830), Westwoodian (1831-1861), Macleayan (1862-1929), Musgravian (1930-1959), and a modern period dominated by professional entomologists.
The early history documents little activity. By the end of the Fabrician period about 60 species had been described, including species described by Linnaeus (1758), and subsequently found in Australia. Fabricius (1775) published the first descriptions of Australian species in the Systema Entomologiae, based on collections made by Joseph Banks during Captain James Cook's expedition to eastern Australia. He described 22 species of bugs, most of which belong to the Pentatomomorpha. In later publications, between 1781 and 1794, Fabricius described 21 additional species. Other European workers, such as Donovan, Thunberg and Kirby, described a handful of Australian species during this period.
Increased taxonomic work during the Westwoodian period resulted in the description of almost 300 more Australian heteropteran species. Between 1837 and 1846 Westwood described about a fifth of the species in this period, the majority in his Catalogue. Erichson (1842) described 21 species of Heteroptera in a review of the fauna of Tasmania. Some 66 Australian pentatomomorphan species were described by Dallas (1851, 1852), based on specimens in the British Museum of Natural History.
Over 1000 Australian heteropteran species were described during the Macleayan period. This period was also characterised by many more workers, including European and Australian taxonomists. It was also the era of the specialised heteropterists, including Carl Stål who, between 1854 and 1876, described 140 species. Other important European works were those of Bergroth (between 1886 and 1927), Reuter (1873 to 1908), and Poppius (1909 to 1921). Walker, in his catalogue of the specimens of Heteroptera in the collection of the British Museum (1867-1873), described 85 species. Between 1880 and 1920, another British worker, W.L. Distant, described almost 150 Australian species in many heteropteran families. The most significant Australian workers of this period were Henry Hacker of the Queensland Museum who described 37 species of Tingidae and one peloridiid in the period from 1927 to 1932, and Herbert Hale of the South Australian Museum who, between 1922 and 1935, described 24 species of aquatic and semiaquatic bugs.
The Musgravian period was characterised by steady taxonomic activity in which some 300 species were added. In this thirty year period, American taxonomists increased their attention to the Australian fauna with Brooks, Hungerford, Drake, Harris, Knight, Scudder, Slater, Todd, Usinger and Wygodzinsky describing almost half of the species. The most significant American worker in the period 1924-1964 was Drake (also with co-authors), who described much of the Australian tingid fauna. Of the European workers, the reduviid taxonomist Miller was a signficant contributor from 1940 to 1959, describing 42 species.
Since the Musgravian Period, over 600 species have been described. Dominant workers include Gross (mostly Pentatomidae), Hill (Dipsocoromorpha), Kormilev (Aradidae), Lansbury (Nepomorpha), Malipatil (Gerromorpha, Reduviidae, Miridae, Lygaeidae), Slater (Lygaeidae, Thaumastocoridae), and Woodward (mostly Lygaeidae), each of whom has produced numerous taxonomic revisions and additions. The major author of Australian Heteroptera in the modern era is G.F. Gross of the South Australian Museum who has described 139 species in the families Anthocoridae, Miridae, Nabidae, Schizopteridae, Berytidae, Lygaeidae, Pentatomidae and Scutelleridae.
For the Coleorrhyncha-Cimicomorpha, the major works since the Musgravian period have been on the Dipsocoromorpha, Gerridae: Halobatinae, some nepomorphans, Miridae and Reduviidae. Between 1980 and 1992, Hill described 61 species of Australian dipsocoromorphans, including the first Dipsocoridae species and many endemic Schizopteridae. Herring (1961) produced a monograph of the marine gerrid genus Halobates Eschscholtz, which preceeded numerous other works on the Halobatinae. Lansbury published more than a dozen papers on the Nepomorpha between 1964 and 1991, predominantly on the Notonectidae, with important revisions of Anisops Spinola and Enithares Spinola (Notonectidae), Sigara Fabricius (Corixidae), and the endemic nepid tribe Goondnomdanepini. The major work on the Australian Miridae in this period was produced by Carvalho & Gross (1982) who described 45 species in a revision of the Phylinae: Leucophoropterini. Wygodzinsky's (1966) monograph of the world Reduviidae: Emesinae provided a comprehensive treatment of the Australian species. In the period 1983 to 1991, Malipatil published major works on the reduviid subfamilies Holoptolinae, Harpactocorinae and the first Australian record of the Visayanocorinae. A more detailed review of taxonomic activity for the Pentatomomorpha will be provided in Volume 27.3B of the Catalogue.
REVIEW OF THE AUSTRALIAN HETEROPTERAN FAUNA
Composition of the Australian Heteropteran Fauna
The Australian Heteroptera comprise 55 families, 779 described genera and 2006 described species (Table 1; Cassis & Gross, Vol. 27.3B, in preparation). Of the families presently represented the Plokiophilidae, Stenocephalidae and Urostylidae are known to occur in Australia but are without described species. A plokiophilid-like cimicoid from northern Queensland is presently being described by Stys (pers. comm.). The Stenocephalidae occur in Australia (Carver et al. 1991) and are represented by an undescribed species of Dicranocephalus Hahn. Carver (et al. 1991) report the occurrence of a urostylid species from northern Queensland and include a habitus figure of a species of Urolabida Westwood. Most of the families not found in Australia are small, often monogeneric, and with restricted, mainly tropical or Western Hemisphere distributions. The Australian fauna includes two endemic families: Hyocephalidae and Lestoniidae.
In Australia, the Coleorrhyncha-Cimicomorpha comprise 352 genera and 909 species (Table 1). This fauna is highly insular with 158 genera (45%) and 757 species (83%) presently recorded as endemic. There is a high level of generic endemicity in the Anthocoridae, Miridae, Notonectidae, Reduviidae, Schizopteridae and Tingidae. Except for Hill's (1980-1992) systematic treatment of the Australian Schizopteridae, the percentage of endemic genera is considered to be exaggerated. The apparently high endemicity for the Anthocoridae, Miridae and Tingidae is probably attributable to paucity of knowledge. Malipatil, in his 1986 to 1991 series of recent revisions of the Reduviidae, culled the numerous monotypic genera described by Miller from 1940 to 1959. The Tingidae require similar revision; the figure of 25 endemic genera may be exaggerated.
Species endemicity in the Coleorrhyncha-Cimicomorpha is at least 50% in all families except the Cimicidae, Omaniidae, Pleidae and Veliidae. For many of the smaller families, including the Aenictopecheidae, Ceratocombidae, Dipsocoridae, Enicocephalidae, Hebridae, Leptopodidae and Schizopteridae, all known species are endemic. Even for the most speciose families, such as the Miridae, Reduviidae and Tingidae, species endemicity is extremely high.
Slater (1982), in an informative review of the Heteroptera, estimated the genus and species numbers for the world fauna. These figures (Table 1) have been amended from the Zoological Record (1980-1984). They indicate that the Australian heteropteran fauna, as presently known, represents about 5% of the world fauna but closer to 10% if the undescribed Miridae are included. Of the Coleorrhyncha-Cimicomorpha families with transoceanic distributions, the Thaumastocoridae, Schizopteridae, Hermatobatidae, Gelastocoridae and Tingidae, are the most taxonomically diverse relative to the world fauna.
The geographical definition of Australia in the Catalogue includes mainland Australia, Tasmania, territorial islands and the Australian Antarctic Territory (see Editorial Preface). Heteroptera have been recorded from Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore Is. and Cartier Is. The number of genera and species in the Coleorrhyncha-Cimicomorpha recognised from these islands are: Christmas Is.-31 genera and 38 species; Cocos (Keeling) Ils-3 genera and 5 species; Lord Howe Is.-26 genera and 30 species; Norfolk Is.-12 genera and 12 species; Ashmore Is. two species; and Cartier Is. one species (Table 3). The Christmas Is. fauna was mostly described by Kirby (1888) and Izzard (1936). Izzard listed 35 species, two of which are now excluded from the fauna. The ubiquitous Nabis capsiformis Germar has been removed from the list, as Izzard's record is probably a misidentification of Nabis kinbergii Reuter (Woodward 1982; Woodward & Strommer 1982). The lygaeid species, Pachybrachius vinctus (Say), is also almost certainly a misidentification as this species is known only from the Western Hemisphere (Slater 1964). Seventeen of the species in Table 3 are known as endemics. No Heteroptera have been recorded from Heard Is., McDonald Is., Macquarie Is. and the Australian Antarctic Territories.
Table 4 lists the number of Coleorrhyncha-Cimicomorpha species found in each Australian state and territory, and in each drainage division. The Queensland fauna presently appears to be the most diverse, with almost two-thirds of the described Australian species represented. Over half of the Australian species are found in the NE coastal drainage division, which is possibly an artifact of the extensive collecting in coastal Queensland. The New South Wales fauna is represented by 316 species, over a third of the Australian fauna, most of which are found in the SE coastal division. The fauna of the Australian Capital Territory is relatively diverse, but this is also an intensively collected area. The faunas of the other mainland states and the Northern Territory are roughly equal in size and relatively depauperate, but this may be partially attributable to limited collecting in Western Australia and the Northern Territory. The Tasmanian fauna comprises 109 species but data from recent collections indicate that many undescribed species exist (Coy et al. 1993).
The Heteroptera exhibit a wide variety of feeding characteristics, including predation, mycetophagy, sap-feeding and mixed-feeding. The host information in the Catalogue refers primarily to feeding records. The majority of families in the Coleorrhyncha-Cimicomorpha are predaceous, except for the Thaumastocoridae, Tingidae and many of the Miridae. There is scant information on host associations for the Australian Heteroptera, particularly for predaceous species. No precise host data are available at present for any species of Coleorrhyncha, Enicocephalomorpha, Dipsocoromorpha, Nepomorpha, Gerromorpha, Leptopodomorpha or Nabidae.
The hosts of very few predatory species of Coleorrhyncha-Cimicomorpha are listed in the Catalogue. Some of these bugs are natural enemies of important economic pests. Orius tantillus (Motschulsky) preys on army worms; Cyrtorhinus lividipennis Reuter and Tytthus chinensis (Stål) prey on delphacid planthoppers on rice and corn; and T. mundulus (Breddin)preys on sugarcane planthoppers and on delphacid planthoppers on rice and corn. The sanguinivorous bedbugs (Cimicidae) and batbugs (Polyctenidae) are often associated with bats; the introduced bedbug, Cimex lectularius L., feeds on a variety of hosts, particularly humans. Of the two polyctenid species, Adroctenes magnus Maa is host specific, whereas Eoctenes intermedius (Speiser) is genus specific and is known from five Taphozous Geoffroy species.
Few host plant records are known for the Australian Coleorrhyncha-Cimicomorpha. Most of the insect/plant associations are obligate. Helopeltis clavifer (Walker) and Campylomma leibknechti (Girault), however, have been recorded on many host species and over a wide variety of plant families.
A number of heteropterists have provided considerable time in reviewing parts of the manuscript. Drs Randall Schuh, Pavel Stys, Nils Möller Andersen, Mary Carver, Mali Malipatil, Jean Péricart and N. Emeljanov provided useful criticisms and additional information. Drs I. Kerzhner and John Polhemus provided exhaustive reviews of many sections of this work. The Catalogue has been improved markedly by their contributions, and the liberal gift of their time and knowledge is especially appreciated.
The two Heteroptera sections of the Catalogue were compiled at the CSIRO Division of Entomology, Canberra and the Australian Museum, Sydney (by G.C.), and at the South Australian Museum, Adelaide (by G.F.G.). The authors wish to thank these institutions for use of their resources and facilities. The authors also wish to thank the Australian Biological Resources Study for their support and commitment to this major project; G.C. was awarded a three year grant and G.F.G a one year grant.
We also wish to thank Ms Gwen Baker and staff of the Information Services Division at the Australian Museum for their support and for obtaining some of the more obscure literature. Administrative support from the Australian Museum greatly facilitated the work and special thanks are due to Dr Des Griffin, Director; Dr Hal Cogger, Deputy Director; and Dr Mike Gray, Head, Invertebrate Division.
Nomenclatural problems were one of the most time consuming aspects of this project. We express our sincere thanks to Dr David McAlpine for his assistance and to other entomologists at the Australian Museum, Dr Dan Bickel, Dr Shane McEvey, Mr Max Moulds and Mr Barry Day, for their patience and encouragement. Ms Sue Lindsay assisted G.C. in many ways, and is especially thanked. At the South Australian Museum, Ms Dzintra Lacis gave many hours of her time and is duly thanked. Also assisting were Ms Debra van Weenen, Ms Maria Drakos and Mrs Kath Ashby.
Many other colleagues were also of great assistance during the preparation of the Catalogue. We especially wish to acknowledge and thank: Dr Paul Arnaud, CAS; Mr Bill Dolling, formerly of the BMNH; Dr Dick Froeschner, USNM; Mr Tom Henry, USDA; Mr Ivor Lansbury, OUM; Dr Jack Lattin, Oregon State University; Mr Gordon Nishida and Dr Scott Miller, BPBM; and Dr Gary Stonedahl, Commonwealth Agricultural Bureau.
The completion of this project would not have been possible without the cooperation of numerous curators of Australian insect collections. We wish to thank: Mr Greg Daniels and Ms Margaret Schneider, EUQ; Mr John Donaldson and Dr Brian Cantrell, QDPI; Dr Murray Fletcher, DARI; Mr Lionel Hill, TDA; Dr Terry Houston, WAM; Dr Mali Malipatil, VAIC; Mr Geoff Thompson, QM; and Dr Ken Walker, NMV. Our sincere thanks also go to Mr Tom Weir, ANIC and Dr Geoff Monteith, QM, for their unfailing assistance and encouragement.
The illustration used in the Aenictopecheidae introduction was drawn by Ms Carmen Zurl, Australian Museum; the Hermatobatidae illustration was kindly provided by Dr Nils M. Andersen and is reproduced with permission from Andersen (1982); the remainder are from Carver et al. (1991) and are reproduced with permission from the CSIRO Division of Entomology and the Melbourne University Press.
Finally, we would like to thank the staff of the Australian Biological Resources Study for their help with this work, especially Drs Keith Houston and Glynn Maynard, whose advice and editorial attention to detail materially improved the Catalogue.
This section of the Zoological Catalogue of Australia database was transferred into the ABRS Platypus software program and edited and prepared for the Australian Fauna Directory by Kathy Tsang and Keith Houston.
The general organisation of this section of the Catalogue
Families, subfamilies, tribes, genera and species are arranged in alphabetical order. This is preferred to a phylogenetic order because of the ease of use of a self-indexing text. The exceptions to alphabetical arrangement are that taxa in incertae sedis and taxa which are unplaced within present subfamilial and tribal classifications are placed at the end of the relevant family, subfamily or tribe.
Family introductions commence with the infraordinal placement of each family and an indication of the number of genera and species worldwide. This is followed by a paragraph on the morphology of the family, which includes a description of the head and appendages, pronotum, metathoracic glands, forewings, legs and genitalia. The biology is described and includes information on habitats and habits, and biological references on Australian species. The suprageneric classification of the family is given, and includes, where applicable, a review of these taxa worldwide, regional distributions, phylogeny, and any alternative taxonomic arrangements. The Australian fauna is summarised and includes the number of genera and species, suprageneric categories, details on the most diverse genera, and significant taxonomic works. Each introduction has its own bibliography and a habitus illustration of a representative species of that family.
Note: Tables and figures contained in the published work (Casis & Gross 1995) and cited here will be included when this section of the Australian Faunal Directory is updated.
In the Catalogue, complete synonymies for genera and species are given, including extralimital synonyms. All junior synonyms have been obtained from the primary literature or catalogues. Where different taxonomic arrangements exist, we have selected the most supported case and list the alternative views, including any putative junior synonyms not accepted in the Catalogue. In most cases, we have given the original authority for each synonymy. Where there is multiple synonymy, each original authority is credited and often the authority of a monographic or revisionary work is also given.
We have maintained strict adherence to the ICZN (1985) as to what constitutes an available name. We do not give incorrect subsequent spellings in the synonymy or the index. They have been listed, in most cases, in the Reference field at the end of each genus or species treatment.
New combinations are given in the Reference field at the end of each species treatment. In most cases, we have given the reference to the authority for the new combination. New combinations can be traced through currently accepted combinations.
Subgeneric and Subspecies Taxonomic Arrangements
The use of subgenera and subspecies in the Heteroptera is common and generally we have followed currently accepted arrangements. The taxonomic authority is given for subgeneric and subspecific arrangements and we list alternative taxonomic arrangements. Type information, extralimital distributions and references are also provided for nominotypical subgenera and subspecies which do not occur in Australia.
We have produced a comprehensive treatment of references that are relevant for the Australian Heteroptera. We have attempted to obtain all primary literature that contains original descriptions or new synonymies. It was often difficult to obtain references for taxa with extralimital synonymies or type localities, and a reference to a secondary source is provided when original sources were not obtained.
A detailed list of references concerned with biology and applied heteropterology has been given for each species in the Reference field. We have not provided complete citations of faunistic works or minor agricultural references. Most citations are accompanied by a brief explanation of the contents in parentheses after the reference and relevant page number(s).
All obtainable information on the primary types (holotypes, syntypes, lectotypes and neotypes) and secondary types (paratypes and paralectotypes) has been provided. This includes reference to any subsequent designation, including any alternative views to a lectotype or neotype designation. The depositories are given for all type specimens and the specimens are designated as seen or not seen (the latter marked by an asterisk). All Australian institutional collections have been examined; in many instances type specimens have been misplaced or lost. We have indicated any discrepancies between the original descriptions and the collections, with qualifications in the Type data field. We have examined specimens held in many of the overseas collections that hold Australian type material, but time and budgetary restrictions prevented a comprehensive treatment of these types. In cases where the type material was not examined, we have used information from secondary sources such as Horn & Kahle (1935-1937) to indicate possible type depositories and such types are marked as not seen. It is probable that numerous corrections will be made to this information by subsequent authors. Where types have been incorporated in collections other than the original depository, we have indicated the current and original placement, e.g. MZUT (ex Cantener coll.). A number of type specimens are listed from personal collections and the whereabouts of some type specimens have not been traced.
For a number of species, we have indicated "holotype" if a single specimen was found in a museum collection and the original description did not indicate more than one type specimen. A number of the early heteropterists, such as Stål, Reuter and Horváth, did not indicate in their original descriptions the number of specimens that the species was described from.
The type localities are given as in the original descriptions, but where label data are different we have given priority to the information found in the literature. We have modernised or anglicised any old Australian localities, e.g. New Holland or NeuHolland is given as Australia, Vandiemensland is given as Tasmania, and Port Denison as Bowen. In such cases, the original citation of the type locality is given in parentheses after the modern locality name. We have also applied this to extralimital type localities, e.g. Ceylon is given as Sri Lanka, Amboina is given as Ambon, and Neu-Guinea as New Guinea. Where the type localities are descriptive and in a foreign language we have provided an anglicised version, e.g. 'collibus Taurinensibus' is given as 'in the hills of Turin, Italy'. Where a type locality is misspelt we have provided a subsequent correction and included the original misspelling in brackets, e.g. '2 miles E of Kiendra' is given as '2 miles E of Kiandra (as Kiendra)'. In a few cases where the type locality could not be found in the Australian gazeteers, we have either made an interpretation of the locality name or left it as originally given, with qualifications.
The Catalogue contains information on the distribution, including extralimital distribution, of genera and species. Family-group distributions are generally provided in the family introductions.
Note: Conversion of the distribution data into the map format will be completed when this section of the Australian Faunal Directory is updated.
Where a genus is not known widely in a region, the distribution of genus groups are given as major zoogeographical regions, within which political descriptors are given, e.g. the distribution of a genus could be given as 'Oriental Region-India, Indonesia, Sri Lanka'. Where the genus is known in more than one country, political descriptors are listed alphabetically under a region. If a genus is widely distributed in zoogeographical regions the distribution is given without political descriptors, such as 'Afrotropical and Oriental Regions', or 'Afrotropical Region-equatorial Africa'.
Species distributions are given firstly as standard drainage divisions or coastal zones (see Map), and then as states, territories and Australian territorial islands. Islands, such as the Torres Strait Ils (QLD), Kangaroo Is. (SA) and Melville Is. (NT), are included under their present state political administration.
The standard drainage divisions are not significant as distributional barriers for Heteroptera but they serve as a convenient means for further subdividing distributions within states and across latitudinal, longitudinal and climatic zones. We have provided point locality information to give the user some indication of the range of a species within a drainage division. This information is given after the standardised descriptors, from which it is separated by a semicolon. Thus a species found widely in the coastal region of NSW could be given as 'SE coastal, NSW; Coffs Harbour, Eden, Lismore, Sydney'. All point localities are listed alphabetically under states or territories. In the Catalogue, many of the records in the drainage divisions, political descriptors and point localities are new and are designated as '(new records)'. These data were obtained by us from museum specimens where identifications were verified. The majority of point localities were obtained from the literature.
Ecological Keywords and Biological Information
A standardised list of ecological descriptors is given in the Editorial Preface. These include life history, feeding and habitat descriptors. Descriptors have been provided for all species in the Catalogue and most species have at least life history and habitat descriptors. We included as much ecological information as possible and assumptions were made liberally. For example, the Anthocoridae are known to be predators of other arthropods and the descriptor 'predator' is given in the Catalogue for all Australian anthocorids, despite little being known on the biology of any of the species. Host information recorded in the literature is given in parentheses in the Ecology field after the 'host' descriptor. In all cases we have indicated the family to which the host belongs. The format used is: host (Plant Name author [Plant family, common name]. The common name refers to that of the plant species or genus. Thus a host record may appear as: host (Eucalyptus L'Hér. [Myrtaceae, gum tree]). The authority for the host information is found in the Reference field and is qualified as a host record. If there is no host indication within the references, the host information can generally be found in the original description of the species.
Other biological information which could not be expressed as an ecological descriptor is given as text after the semicolon at the end of the descriptors. This information includes habitat information, e.g. larvae and adults found in and under decaying vegetation of Pandanus [Pandanaceae].
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.
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
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.
Coy, R., Greenslade, P. & Rounsevell, D. 1993. A Survey of Invertebrates in Tasmanian Rainforests. Hobart : Tasmanian Parks and Wildlife Service. National Rainforest Conservation Program Report No. 9 xi 104 pp.
Distant, W.L. 1902. The Fauna of British India, including Ceylon and Burma. Rhynchota. (Heteroptera). London : Taylor & Francis Vol. 1 pp. xxviii 1-432. [1902: vol. 1 438 pp.; 1903–1904: vol. 2 503 pp.; 1906: vol. 3 503 pp.; 1908: vol. 4 501 pp.; 1910: vol. 5 362 pp.; 1918: vol. 6 210 pp.]
Evans, J.W. 1981. A review of present knowledge of the family Peloridiidae and new genera and new species from New Zealand and New Caledonia (Hemiptera: Insecta). Records of the Australian Museum 34: 381-406
Fabricius, J.C. 1775. Systema Entomologiae, sistens Insectorum Classes, Ordines, Genera, Species, adiectis Synonymis, Locis, Descriptionibus, Observationibus. Flensburgi et Lipsiae [= Flensburg & Leipzig] : Kortii xxxii 832 pp. [Date published 17 April]
Hacker, H. 1929. New species of Australian Tingitidae (Hemiptera). Memoirs of the Queensland Museum 9: 324-334 pls XXXII-XXXV
Horn, W. & Kahle, I. 1937. Über entomologische Sammlungen (Ein Beitrag zur Geschichte der Entomo-Museologie). Berlin : Dahlem W. Horn. [1935: vol. 1 vi pp. 1–160 pls I–XVI; 1936: vol. 2 pp. 161–296 pls XVII–XXVI; 1937: vol. 3 pp. 297–536 pls XXVII–XXXVIII]
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.
Kristensen, N.P. 1991. Phylogeny of extant hexapods. pp. 125-140 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.
Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. pp. 141-179 in CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. Melbourne : Melbourne University Press Vol. 1 xiii 542 pp.
La Rivers, I. 1974. Catalogue of taxa described in the family Naucoridae (Hemiptera) supplement no. 1: corrections, emendations and additions, with descriptions of new species. Memoirs. Biological Society of Nevada, Occasional Papers 38: 1-17
Lethierry, L. & Severin, G. 1896. Catalogue Général des Hémiptères. Tome III. Hétéroptères Tingidae, Phymatidae, Aradidae, Hebridae, Hydrometridae, Henicocephalidae, Reduvidae, Saldidae, Apophilidae, Ceratocombidae, Cimicidae, Anthocoridae. Berlin : R. Friedländer & Fils 275 pp.
Linnaeus, C. 1758. Systema Naturae per Regna tria Naturae, secundem Classes, Ordines, Genera, Species, cum Characteribus, Differentis, Synonymis, Locis. Tom.1 Editio decima, reformata. Holmiae : Laurentii Salvii 824 pp.
Lundblad, O. 1933. Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von Sumatra, Java und Bali auf Grund des Materials der Deutschen Limnologischen Sunda-Expedition, nebst Revision einiger anderer, indoaustralischer Arten. Archiv für Hydrobiologie 12: 1-195, 263-489
Mahner, M. 1993. Systema phylogeneticum Cryptoceratorum (Insecta, Heteroptera). Zoologica (New York) Stuttgart 48(143): ix + 302 pp.
Myers, J.G. & China, W.E. 1929. The systematic position of the Peloridiidae as elucidated by a further study of the external anatomy of Hemiodoecus leai China. Annals and Magazine of Natural History 3: 282-294
Oshanin, B.F. 1906. Verzeichnis der Palaearktischen Hemipteren mit besondere Berücksichtigung ihrer Verteilung im Russischen Reiche. Band I Heteroptera. Supplement to Annuaire du Musée Zoologique de l'Académie Imperiale des Sciences, St Petersburg. pp. i–lxxiv, 1–393.
Oshanin, B.F. 1910. Verzeichnis der Palaearktischen Hemipteren mit besondere Berücksichtigung ihrer Verteilung im Russischen Reiche. Band III Nachträge und Verbesserungen zum I und II, Bande (Ende des Werkes). Supplement to Annuaire du Musée Zoologique de l'Académie Imperiale des Sciences, St Petersburg. pp. 587–1087
Polhemus, J.T. 1979. Family Naucoridae—Creeping Water Bugs, Saucer Bugs. pp. 131-138 in Menke, A.S. (ed.). The Semiaquatic and Aquatic Hemiptera of California (Heteroptera: Hemiptera). Berkeley : University of California Press.
Schlee, D. 1969. Morphologie und Symbiose; ihre Beweiskraft für die Verwandtschaftsbeziehungen der Coleorrhyncha (Insecta, Hemiptera). Phylogenetische Studien an Hemiptera IV: Heteropteroidea (Heteroptera + Coleorrhyncha) als monophyletische Gruppe. Stuttgarter Beiträge zur Naturkunde 210: 1-27
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
Slater, J.A. 1964. Hemiptera (Heteroptera): Lygaeidae. pp. 15-228 in Hanström, B., Brinck, P. & Rudebeck, G. (eds). South African Animal Life. Results of the Lund University Expedition in 1950–1951. Stockholm : Almqvist & Wiksell Vol. 10.
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]
Štys, P. 1989. [Phylogenetic systematics of the most primitive true bugs (Heteroptera, Enicocephalomorpha, Dipsocoromorpha).]. Práce Slovenská entomologická spolocnost' SAV, Bratislava 8: 69-85 [In Czech]
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. & Strommer, N. 1982. Nabis kinbergii Reuter, the current name for Tropiconabis nigrolineatus (Distant), and its Australian distribution (Hemiptera: Nabidae). Journal of the Australian Entomological Society 21: 306
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