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Jewel Bugs of (Insecta, , )1

G. CASSIS & L. VANAGS

Abstract: The Australian genera of the Scutelleridae are redescribed, with a exemplar of the ma- le genitalia of each illustrated. Scanning electron micrographs are also provided for key non-ge- nitalic characters. The Australian jewel bug fauna comprises 13 genera and 25 species. Heissiphara is described as a new genus, for a single species, H. minuta nov.sp., from . Calliscyta is restored as a valid genus, and removed from synonymy with Choerocoris. All the Australian species of Scutelleridae are described, and an identification key is given. Two new species of Choerocoris are des- cribed from eastern Australia: C. grossi nov.sp. and C. lattini nov.sp. Lampromicra aerea (DISTANT) is res- tored as a valid species, and removed from synonymy with L. senator (FABRICIUS). nobilis (LIN- NAEUS) is recorded from Australia for the first time. Calliphara billardierii (FABRICIUS) and C. praslinia praslinia BREDDIN are removed from the Australian biota. The identity of Sphaerocoris subnotatus WAL- KER is unknown and is . A description is also given for the Neotropical species, Agonoso- ma trilineatum (FABRICIUS); a biological control agent recently introduced into Australia to control the pasture weed Bellyache Bush ( gossypifolia, ). Coleotichus borealis DISTANT and C. (Epicoleotichus) schultzei TAUEBER are synonymised with C. excellens (WALKER). Callidea erythrina WAL- KER is synonymized with Lampromicra senator. Lectotype designations are given for the following taxa: Coleotichus testaceus WALKER, Coleotichus excellens, Sphaerocoris circuliferus (WALKER), Callidea aureocinc- ta WALKER, Callidea collaris WALKER and Callidea curtula WALKER. A discussion of the higher classifica- tion of scutellerids is given, and diagnoses of the subfamilies natively present in Australia are provided. A new term, the ejaculatory apparatus is introduced for the subdistal elaboration of the ductus seminis, to include the ventral conducting canal, ejaculatory reservoir and dorsal conducting canal. A survey of male pheromonal glands found on the pregenital abdominal sterna was undertaken, recognising two ty- pes, androconial glands (present in Odontotarsinae and Tectocorinae) and setose sternal glands (Elvi- surinae). The biology of the Australian species is summarised, with new host records given for Austrotichus rugosus GROSS, all the Australia native species of Coleotichus (C. artensis (MONTROUZIER), C. costatus FABRICIUS and C. excellens), Calliscyta stalii (VOLENHOVEN), Scutiphora pedicellata (KIRBY) and (HAHN).

Key words: androconial glands, Australia, biogeography, classification, ejaculatory apparatus, Heterop- tera, host , new genus, new species, , Scutelleridae, setose sternal glands.

Introduction ocorium of the forewings minimally exposed proximally, and the membrane tip some- The Scutelleridae or jewel bugs are a times visible caudally (e.g. Scutellerinae: of landbugs belonging to the super- Calliphara, Lampromicra). Scutellerids can be family Pentatomoidea (Insecta, Heteropte- confused with other pentatomoids with an ra) (SLATER 1982; SCHAEFER 1993; SCHUH & enlarged scutellum, including members of SLATER 1995; CASSIS & GROSS 2002). These the cydnid complex (e.g. Corimelaeninae), often large are recognised in part by Aphylidae, Canopidae, Lestoniidae, Platasp- the greatly enlarged scutellum, which covers idae, and a number of genera of the nomino- most if not all of the abdomen, with the ex- typical subfamily of the (e.g.

1 This work is dedicated to our friend and colleague Dr Ernst Heiss, in recognition of his efforts in documenting true Denisia 19, zugleich Kataloge bugs, particularly piesmatids and barkbugs from around the world. In celebrating his 70th birthday, we dedicate a new der OÖ. Landesmuseen species, Heissiphara minuta, in his honour; the shortest described jewel bug for the tallest heteropterist! Neue Serie 50 (2006), 275–398

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Australian genera: Ippatha DISTANT, Kam- and corrections. Illustrations are given for a butha DISTANT). Most species of the species of each genus, and for each species a nominotypical subfamily, the Scutellerinae, detailed description is given, and for those and some taxa of the Pachycorinae and taxa without illustrations in this work, users Sphaerocorinae, are characterised by highly are referred to those given in MCDONALD colourful bodies, presumably aposematic. In (1961, 1963a, 1963b) and MCDONALD & contrast, the adults of species of other scutel- CASSIS (1984). The biological control lerid subfamilies are often dull in coloura- agent, Agonosoma trilineatum (FABRICIUS) tion, and apparently cryptozoic (e.g. Elvi- (Scutelleridae: Pachycorinae), introduced surinae, Eurygastrinae and Odontotarsinae), from to control a pasture although in some such cases the larvae are weed (Bellyache Bush, Jatropha gossypifolia, brilliantly coloured (e.g. Elvisurinae: Euphorbiaceae) (HEARD et al. 2002), is also Coleotichus). treated in this work. In addition, we review the status of scutellerid systematics and Jewel bugs are moderately diverse in character systems, place the Australian comparison to other pentatomoid families jewel bug fauna in a global context, and dis- (e.g. : 180 species; Cyd- cuss their biology. nidae: 600; Pentatomidae: 6000; Plataspi- dae: 560; and, : 235; figures from CASSIS & GROSS 2002). LATTIN Material and Methods (1964) recognised 80 genera and 450 species of jewel bugs worldwide. In the interim, less Specimens than 50 new taxa have been described Most of the material examined includes (JAVAHERY et al. 2000; CASSIS & GROSS new collections across temperate Australia 2002), countered by significant levels of by the senior author [GC] and Randall synonymy (e.g. EGER & LATTIN 1995). Schuh of the American Museum of (AMNH). Additional material was FABRICIUS (1775) described the first examined from the Museum Australian scutellerid species, for the charis- (QM), the Australian National Col- matic jewel bug species Calliphara imperialis, lection (ANIC), the South Australian Mu- Calliphara regalis and Choerocoris paganus. seum (SAMA), and the Western Australian STÅL (1873) produced the first comprehen- Museum (WAM). Late in the project, one sive and systematic treatment of scutel- of us [GC] was able to re-examine type and lerids, providing a suprageneric classifica- other materials in the Natural History Mu- tion, including keys to the tribes and genera, seum (BMNH). We particularly focused on as well as listing all described species, in- the Dallas, Distant and Walker types, doubt- cluding those from Australia. SCHOUTEDEN ing the validity of previous observations on (1904) provided a monograph of the jewel their status. The type material that we ex- bugs, including detailed descriptions of the amined is listed separately, and new lecto- supraspecific taxa and a list of species. The type and paralectotype designations are giv- Australian scutellerid fauna has received a en. The Fabricius types in the Banks collec- number of modern treatments, including tion (BMNH) were not available for obser- those of GROSS (1975) and MCDONALD & vation. We also found that some of the lec- CASSIS (1984). CASSIS &GROSS (2002) cat- totype designations of LYAL (1979) for Cal- alogued the Australian scutellerids, recog- liphara are invalid. For types that we have nizing 11 genera and 26 species. In this work not re-examined, we refer users to CASSIS & we give an updated revision of the Aus- GROSS (2002), which lists all the types and tralian scutellerids, providing generic and associated information; most of which have species treatments, including the descrip- been previously examined by one of us tion of new taxa and addition of new char- [GC]. acter information. An identification key is given for all genera and species. This work The distribution maps are based on these serves as an updated account of MCDONALD new data as well as the collection localities & CASSIS (1984) and CASSIS & GROSS given in GROSS (1975), MCDONALD & CAS- (2002), and includes numerous additions SIS (1984) and CASSIS & GROSS (2002).

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Microscopy es of second conjunctival appendages; Observations were made using Leica MZ CAII(L) = lateral branch of first conjuncti- 16A and MZ 12 stereomicroscopes, and a val appendages; CAIII = paired or fused Leica DMB compound microscope. Illustra- third conjunctival appendage(s); CAIII(D) tions of morphological features were made = dorsal branch of third conjunctival ap- using a camera lucida. These were rendered pendages; and, CAIII(V) = ventral branch in Adobe Illustrator software. Scanning of third conjunctival appendages. Further electron micrographs were made using a abbreviations are provided in the illustra- Cambridge scanning microscope. tions, where legends are provided for each plate. For Heissiphara minuta nov.sp. and Morphometric measurements Calliphara nobilis, the following abbrevia- tions are also used for the morphometric The measurements taken in MCDONALD characters: BL = body length; IOD = inte- & CASSIS (1984) are recorded for the new rocular distance; and, PW = pronotal width. species (Choerocoris grossi nov.sp., C. lattini nov.sp. and Heissiphara minuta nov.sp.) or Biology species of revised status (Lampromicra aerea In Australia, the biology of only a few DISTANT). Measurements are also given for scutellerid species has been described in any Austrotichus rugosus GROSS, which were not detail. MCDONALD (1960, 1963b) studied given in MCDONALD & CASSIS (1984), and the introduced species Agonosoma trilinea- the life cycle, larval morphology and feeding tum. All measurements are given in tables, behaviour of five Australian species (Lam- promicra senator parentum Scutipho- except for Heissiphara minuta nov.sp., for , , ra pedicellata, Choerocoris paganus and Tecto- which only the holotype was measured, and coris diophthalmus). The biology of Tectocoris the new Australian record, Calliphara nobilis diophthalmus has also been detailed, often in (LINNAEUS); in both cases, measurements relation to its status (BALLARD & are given in the species description. Mea- HOLDAWAY 1926; MCKEOWN 1942; STAD- surements were made using a graticule, and DON et al. 1987). CASSIS & GROSS (2002) all measurements are maximum lengths and gave a brief summary of the biology of Aus- given in millimetres. tralian scutellerids and a list of their host In MCDONALD & CASSIS (1984), the plants. The following review focuses on the antennae are referred to as five-segmented; plant associations and feeding behaviour of in fact, they are four-segmented with AII these species. subdivided (see below). The following syn- onymy is required to allow for comparison of Plant Associations measurements in the latter work, and those From a systematic heteropterist’s point given in below: AII = AII(a); AIII = AII(b); of view, the fundamental biological infor- AIV = AIII; and, AV = AIV. mation source for phytophagous species is the documentation of plant associations. In Morphological Abbreviations this work we have increased the number of Throughout this manuscript abbrevia- associations by almost 50 % from those giv- tions are used for antennal and labium seg- en in CASSIS & GROSS (2002), with our own ments, pregenital abdominal sterna, and the fieldwork resulting in 10 new plant associa- conjunctival appendages of the male aedea- tion records (Table 1). Of the 25 species of gus, as follows: AI-AIV = first to fourth an- Australian scutellerids (including Agonoso- tennal segments, with AII divided, and giv- ma trilineatum), 16 species have known en as AII(a) and AII(b); LI-LIV = first to plant associations. In Table 1 we list all the fourth labium segment; SII-SVIII = first to known plant associations, and report or eighth abdominal sterna; CAI = first con- speculate as to how they use these plants. junctival appendages; CAI(D) = dorsal CASSIS et al. (in press) have shown in Aus- branch of first conjunctival appendages; tralian , that many fami- CAI(V) = ventral branch of first conjuncti- ly-groups have a broad range of plant associ- val appendages; CAII = second conjuncti- ations, including scutellerids (CASSIS & val appendages; CAII(M) = medial branch- GROSS 2002). As with other pentatomoids,

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Table 1: Plant associations (species, and ) for Australian jewel bugs, and information source. CASSIS & GROSS (2002) includes other citations to host plant information. GC = G. CASSIS fieldwork 1995-2005.

Jewel Bug Species Plant Species Association Plant Family Plant Order Clade Plant Usage Source Austrotichus rugosus humilis Eurosids I food plant this work; GC fieldwork (OTTO & F.DIETR.) L.A.S.JOHNSON Casuarinaceae Fagales Eurosids I food plant this work; GC fieldwork Allocasuarina pusilla (MACKLIN) L.A.S.JOHNSON Coleotichus artensis excelsa Eurosids I unknown this work; label data (FENZL) BENTH. Coleotichus costatus consanguinea Mimosaceae Eurosids I food plant this work; GC fieldwork R.S.COWAN & MASLIN Acacia cyclops G.DON Mimosaceae Fabales Eurosids I food plant CASSIS & GROSS 2002 Acacia ligulata BENTH. Mimosaceae Fabales Eurosids I food plant WHITNEY & STANTON 2004 Acacia pendula A.CUNN EX G.DON Mimosaceae Fabales Eurosids I food plant CASSIS & GROSS 2002 Acacia saligna (LABILL.) H.L.WENDL. Mimosaceae Fabales Eurosids I food plant CASSIS & GROSS 2002 Acacia stowardii MAIDEN Mimosaceae Fabales Eurosids I food plant this work; GC fieldwork Acacia victoriae BENTH. Mimosaceae Fabales Eurosids I food plant CASSIS & GROSS 2002 viscosa JACQ. Fabales Eurosids I food plant Whitney pers. comm. inermis R.BR. Core Eudicot ?sitting record this work; GC fieldwork Coleotichus excellens Ficus sp. Moraceae Rosales Eurosids I ?food plant this work; GC fieldwork Hibiscus rosasinensis L. Eurosids II unknown this work; label data Solenotichus circuliferus gongylocarpa BLAKELY ?aggregation CASSIS & GROSS 2002 (unplaced) site Agonosoma trilineatum Jatropha gossypifolia L. Euphorbiaceae Eurosids I food plant Heard pers. comm. Calliphara nobilis Acer oblongum WALL. Sapindaceae Eurosids II food plant TAKARA 1957 Xanthium strumarium L. Asteraceae Asterales Euasterids II food plant BALOCH et al. 1968 tanarius (L.) MULL.ARG. Euphorbiaceae Malpighiales Eurosids I food plant HILGENDORF & GADEN 1982 agallocha L. Euphorbiaceae Malpighiales Eurosids I food plant LIM et al. 2001 Rhizophora sp. Rhizophoraceae Malpighiales Eurosids I food plant LIM et al. 2001 Calliscyta stalii Pandanus sp. Pandanaceae Pandanales Monocots ?aggregation Monteith pers. comm. site Cantao parentum claoxyloides (F.MUELL.) MULL.ARG. Euphorbiaceae Malpighiales Eurosids I food plant CASSIS & GROSS 2002 (LAM.) MULL.ARG. Euphorbiaceae Malpighiales Eurosids I food plant CASSIS & GROSS 2002 Mallotus discolor BENTH. Euphorbiaceae Malpighiales Eurosids I food plant Monteith pers. comm. Araucariaceae Coniferales Gymnosperms aggregation CASSIS & GROSS 2002 AITON EX A.CUNN. site Choerocoris grossi viscosa (LABILL.)MIQ. Euphorbiaceae Malpighiales Eurosids I ?food plant label data Choerocoris paganus Dodonaea lanceolata F.M UELL. Sapindaceae Sapindales Eurosids II food plant CASSIS & GROSS 2002 Dodonaea triquetra J.C.WENDL. Sapindaceae Sapindales Eurosids II food plant CASSIS & GROSS 2002 Sapindaceae Sapindales Eurosids II food plant CASSIS & GROSS 2002 Dodonaea viscosa angustissima (DC.) J.G. WEST Sapindaceae Sapindales Eurosids II food plant this work; GC fieldwork Dodonaea viscosa mucronata J.G.WEST Sapindaceae Sapindales Eurosids II food plant this work; GC fieldwork Dodonaea viscosa spatulata (SM.) J.G. WEST Sapindaceae Sapindales Eurosids II food plant this work; GC fieldwork Medicago sativa L. Fabales Eurosids I ?food plant CASSIS & GROSS 2002 Plantago sp. Plantaginaceae Lamiales Euasterids I ?food plant CASSIS & GROSS 2002 Choerocoris variegatus (DC.) BAILL. Euphorbiaceae Malpighiale Eurosids I food plant CASSIS & GROSS 2002 (LABILL.) MIQ. Euphorbiaceae Malpighiale Eurosids I food plant CASSIS & GROSS 2002 Spyridium globulosum (LABILL.) BENTH. Rhamnaceaes Rosales Eurosids I food plant CASSIS & GROSS 2002 Acacia cyclops G. DON. Mimosaceae Fabales Eurosids I ?sitting label data; record GC fieldwork Heissiphara minuta Stenanthemum complicatum Rhamnaceae Rosales Eurosids I ?food plant label data; GC fieldwork (F. MUELL.) RYE Lampromicra aerea spinosa CAV. Euasterids II unknown this work; label data ambigua (SM.) DRUCE Myrtaceae Myrtales Rosids (unplaced) unknown this work; label data Lampromicra senator Adiantum aethiopicum L. Adiantaceae Filicales Ferns ?aggregation this work; GC fieldwork site Bursaria sp. Pittosporaceae Apiales Euasterids II food plant this work; label data Arachis hypogaea L. Fabaceae Fabales Eurosids I ?aggregation CASSIS & GROSS 2002 site oblongifolia (MULL.ARG.) MULL. ARG. Euphorbiaceae Malpighiales Eurosids I food plant CASSIS & GROSS 2002

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Jewel Bug Species Plant Species Association Plant Family Plant Order Clade Plant Usage Source Lampromicra senator Ficus carica L. Moraceae Rosales Eurosids I food plant CASSIS & GROSS 2002 Ficus leucotricha (MIQ.) MIQ. Moraceae Rosales Eurosids I food plant CASSIS & GROSS 2002 Lantana camara L. Verbenaceae Lamiales Euasterids I unknown CASSIS & GROSS 2002 Mimosa pigra L. Mimosaceae Fabales Eurosids I unknown WILSON et al. 1990 Scutiphora pedicellata Crataegus oxyacantha L. Rosaceae Rosales Eurosids I food plant CASSIS & GROSS 2002 Dodonaea triquetra J.C.WENDL. Sapindaceae Sapindales Eurosids I food plant CASSIS & GROSS 2002 Prunus domestica L. Rosaceae Rosales Eurosids I food plant CASSIS & GROSS 2002 Sambucus guadichaudiana DC. Caprifoliaceae Dipsacales Euasterids II food plant http://morwell np.pangean.net Ficus sp. Moraceae Rosales Eurosids I food plant FROGGATT 1907 Leptospermum sp. Myrtaceae Myrtales Rosids (unplaced)?sitting record label data

Tectocoris diophthalmus Gossypium hirsutum L. Malvaceae Malvales Eurosids II food plant CASSIS & GROSS 2002 Hibiscus tiliaceus L. Malvaceae Malvales Eurosids II food plant CASSIS & GROSS 2002 patersonia (ANDREWS) G.DON Malvaceae Malvales Eurosids II food plant this work; GC fieldwork acerifolius (A.CUNN EX G.DON) MACARTHUR & C.MOORE Malvaceae Malvales Eurosids II food plant Monteith pers. comm. (SCHOTT & ENDL.) TERRACINO Malvaceae Malvales Eurosids II food plant Monteith pers. comm. (T.MITCH EX. LINDL.) K.SCHUM. Malvaceae Malvales Eurosids II food plant Monteith pers. comm. scutellerids generally prefer higher an- where in the Heteroptera (e.g., SLATER giosperms (sensu STEVENS 2006; Angio- 1971; CASSIS & GROSS 2002; Heterogastri- sperm Phylogeny Group and clas- dae), including extralimital scutellerids sification), belonging to the rosid and aster- (CERVANTES 2004). id subclasses, but particularly with species The association of Australian scutel- belonging to three groups of the rosids: i. lerids with asterid angiosperms is far less Unplaced Rosids; ii. Eurosid I; and, iii. Eu- , with records known for Euasterid I rosid II. More than half of all Australian and Euasterid II plants. The associations scutellerids have been found on ‘Eurosid I’ plants, with highest frequency of association with more angiosperms are not of any with the Euphorbiaceae, Sapindaceae and great significance, with the association with Mimosaceae. Australian scutellerids are ‘Core Eudicot’ plants most likely ‘sitting known from a range of euphorbs, with three records’ rather than host plants. Likewise, species (Cantao parentum, Choerocoris pa- the association of the above scutellerids ganus and Lampromicra senator) found in with gymnosperms and ferns are considered Australia on three different euphorb genera, either sitting or aggregation records. and Calliphara nobilis found on two euphorb Host Plant Specificity and Food species outside of Australia (). Preferences The sapind records are of particular interest, with a number of species found in associa- Unlike the , where host plant tion with species of Dodonaea, particularly specificity is very high (SCHUH & SLATER D. viscosa and D. triquetra. Choerocoris pa- 1995; CASSIS et al. in press), the Australian ganus, which has a broad range of plant as- scutellerid species are found on a broad sociations, is most commonly found on range of plants. For example, only three jew- Dodonaea viscosa (including a number of its el bug species are known from one plant subspecies) (Fig. 1d). This plant is a second- species alone (Table 1: Coleotichus artensis, ary host plant for Coleotichus costatus (Whit- Solenotichus circuliferus and Calliscyta stalii), ney pers. comm.), which is most commonly but in each of these cases, the species are found on a range of Acacia species. Curious- not commonly encountered, and there is ly, this ‘Acacia and Dodonaea’ pattern of as- doubt as to whether any of the records rep- sociation also occurs for the Hawaiian en- resent food or breeding hosts. For scutel- demic Coleotichus species, C. blackburniae lerids where more information exists, Can- (JOHNSON et al. 2004). Three species of tao parentum is found on three species of the scutellerids (Coleotichus excellens, Lampromi- euphorb genus Mallotus (Fig. 1b). This is al- cra senator and Scutiphora pedicellata) are so the case for Austrotichus rugosus, which found on figs; an association known else- we have found on two species of Allocasuar-

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ina (Casuarinaceae) in temperate Australia. species of the Australian endemic genus Tectocoris diophthalmus also shows a degree of Morbora have only been collected on the host plant restrictiveness, as it is found only ground (see below). In contrast, Tecotocoris on species of plants belonging to the Mal- diophthalmus feeds on vegetative and repro- vaceae. ductive parts of their host plants (BALLARD & HOLDAWAY 1926; MCDONALD 1963c). Scutellerids are considered to be exclu- sively phytophagous although a few cases of Bellyache plant, Jatropha gossypifolia, is a omnivory and opportunistic carnivory are native of the Neotropical region, and is be- known (EUBANKS et al. 2003). Surprisingly coming a significant pasture weed in north- little is known of the biology of jewel bugs, ern Australia (HEARD et al. 2002; Heard aside from the economic Sunn pests (JAVA- pers. comm.). In 2003, the South American HERY et al. 2000; integriceps and scutellerid species Agonosoma trilineatum, related species). In Australia, Tectocoris dio- was introduced into southeast Queensland phthalmus has been recorded as a minor pest as a biological control agent. This species of cotton, where it is known to facilitate has not as yet been recovered in Australia. staining of the bolls (BALLARD 1925, 1927; BALLARD & HOLDAWAY 1926). Three Aggregation and Maternal Behaviour species (Cantao parentum, Lampromicra sena- Numerous species of scutellerids are tor and Scutiphora pedicellata) have been re- known to exhibit aggregative behaviour. ported as minor pests of cherries (FROGGATT JAVAHERY et al. (2000) report that Eurygaster 1901, 1902, 1907; MCDONALD 1963c), al- integriceps is found in mass aggregations in though there are no recent records of pest the Palearctic region; it both aestivates in activity. the mid-summer and overwinters in the As with many pentatomomorphans, nu- winter in such aggregations. MONTEITH merous scutellerids are known to feed on (1982) found Lampromicra senator in dry season aggregations amongst vegetation and . JAVAHERY et al. (2000) report that some species also feed on , as well as on the ground in tropical north Queens- vegetative parts. The Australian species land. We have also recently found this feed on pre-dispersed seeds, upon their host species at the beginning of the dry season in plants. They are also commonly encoun- large aggregations on fig in open tered on the ground, in the ‘ shadow’ of woodland on the Atherton Tableland. MC- their host plant, feeding also on post-disper- DONALD (1963c) reports that Scutiphora sal seeds. The Australian species that have pedicellata hibernates under the bark of Eu- been encountered on the ground as well as calyptus , and Cantao parentum hiber- on plants include, Austrotichus rugosus, nates in clusters on the underside of leaves. Coleotichus costatus, Choerocoris paganus, C. Many of the small clusters of Cantao paren- variegatus, Lampromicra senator and Scutipho- tum merge as winter progresses leading to the formation of giant aggregations of some- ra pedicellata. MCDONALD (1963c) reported Lampromicra senator feeding on the fruits of times tens of thousands of bugs in the tops of Breynia . There is also strong evidence emergent trees (often Hoop pines, Araucaria that a number of Odontotarsinae species are cunninghamii) adjacent to the thickets of epigaeic; found on the ground, presumably their Mallotus food plants (Monteith pers. being post-dispersal seed-predators. All the comm.). Monteith (pers. comm.) has also found Calliphara regalis in aggregations on surfaces (Fig. 1a).

Fig. 1: Examples of Australian Scutelleridae Scutellerids are often found in large ag- species and their host plant species gregations when feeding (e.g., Choerocoris (a) adult aggregation of Calliphara regalis paganus), and the larvae are commonly asso- (b) Cantao parentum on Mallotus philippensis (c) Choerocoris paganus on ciated with the adults. Some species are also Dodonaea viscoca (d) Lampromicra senator known to engage in maternal care, with fe- on unidentified host plant (e) Scutiphora males protecting the egg brood from preda- pedicellata on unidentified host plant (f) Tectocoris diophthalmus on tors and parasites. In Australia, this has Brachychiton acerifolius. been recorded for Tectocoris diophthalmus

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Table 2: Alternative Suprageneric Classifications of the Scutelleridae.

DALLAS (1851) STÅL (1872, SCHOUTEDEN KIRKALDY (1909) LESTON (1952d) MCDONALD GROSS (1975) MCDONALD & CARAYON CASSIS & 1873) (1904) (1966) CASSIS (1984) (1984) GROSS 2002 Superfamily? Subfamily Subfamily Subfamily Subfamily Family Family Family Family Family Scutelleroides Scutellerinae Scutellerinae Scutellerinae Scutellerinae Scutellerinae Scutelleridae Scutelleridae Scutelleridae Scutelleridae Pachycoridae Elvisuraria Elvisuraria Elvisurini Eurygastrini: Eurygastrinae Elvisurinae Elvisurinae Eurygastrinae: Elvisurinae Eurygastraria Eurygastrini Eurygastridae Eurygastraria Odonto- Odontotarsini Eurygastrini: Odonto- Scutellerinae Odonto- Eurygastrinae: Odonto- tarsaria Odontoscelaria scelinae tarsinae Odontoscelini tarsinae Odonto- Odonto- Scutelleraria Scutellerini Eurygastrini: PachycorinaeOdonto- Scutellerinae Eurygastrinae: Pachy- scelidae scelaria Odontotarsaria tarsinae Odontotarsini corinae Odonto- Sphaero- Sphaerocorini Pachycorini Scutellerinae Tectocorinae Pachycorinae Scutellerinae tarsaria coraria Scutelleraria Tetyraria Tetyrini Scutellerini: Scutellerinae: Tectocorinae Elvisuraria Elvisurini Sphaero- Scutellerini: Scutellerinae: coraria Scutelleraria Scutellerini Tetyraria Scutellerini: Scutellerinae: Sphaerocoraria Sphaerocorini

where the eggs are laid on petioles and the been surveyed extensively across the family. females guard the eggs (DODD 1904; TAL- As with other taxa in the Pentatomoidea LAMY & SCHAEFER 1997; TALLAMY 1998; the jewel bugs are polythetically defined. MONTEITH 2006). KUMAR (1965) and GAFFOUR-BENSEBBANE (1990) gave features of the male genitalia Higher classification (‘six ectadene tubules’) and spermatozoa (‘plumed’ motility structure) as scutellerid and Supra-familial characters, but the sampling was limited. position of Scutelleridae within the Pentatomoidea GAPUD (1991), in the first phylogenetic analysis of pentatomoid relationships, pro- LEACH (1815) first recognised the jewel posed that scutellerids were diagnosable by bugs as a family-group. Subsequent workers the following characters: i. weak to moder- proposed alternative classifications, with ately sulcate prosternum; ii. ten or more scutellerids as a family (e,g., VAN DUZEE forewing membrane veins; iii. second gono- 1917) allied to the Pentatomidae, or as a coxae fused; iv. first gonapophyses membra- subfamily of the Pentatomidae, along with neous, with first rami reduced; and, v. sec- other pentatomoid taxa such as the Cyd- ond gonapophyses membraneous, with sec- nidae, and Tessaratomidae (e.g. ond rami weakly sclerotized; and, vi. sper- KIRKALDY 1909; LESTON 1952c, 1952d). In matheca with sclerotized basal groove. the modern era, scutellerids are consistently SCHUH & SLATER (1995) and CASSIS & maintained as a distinct family (CARVER et GROSS (2002) provided broader diagnoses of al. 1991; SCHAEFER 1993; SCHUH & SLATER the family, which repeated the aforemen- 1995; CASSIS & GROSS 2002). At present, tioned characters, and also highlighted the the debate centres on the systematic place- following: i. absence or reduction of the fre- ment of the Scutelleridae in the Pentato- na; ii. sulcate prosternum; and, iii. carinate moidea, for example: i. sister-taxon to the propleura. GRAZIA et al. (in prep.) have Canopidae, basal to the ((Lestoniidae + identified these latter characters as diagnos- Plataspidae) + (Thyrecorinae + )) tic, but not exclusive to the Scutelleridae. (GAPUD 1991); ii. Scutelleridae + Tessarato- They report that the grooved spermathecal midae, basal to the Pentatomidae (DAI & base, first identified by GAPUD (1991), is ex- ZHENG 2004); iii. Scutelleridae + Penatomi- clusive for the Scutelleridae; curiously they dae or (Pentatomidae + (Scutelleridae + do not include it in their phylogenetic (Tessaratomidae + ) (FISCHER analysis. FISCHER (2001) proposed that the 2001); and, iv. sister-taxon to the Thaumas- scutellerids are defined by the following tellidae (GRAZIA et al. in prep.). characters: i. greatly enlarged scutellum; ii. The Scutelleridae are not currently di- first abdominal glands laterally positioned; agnosed by any single character that has iii. hindwing with a hamus; and, iv. the cori-

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um of the forewing desclerotized. He also lowed the classification of Schouteden, and Fig. 2: Abdominal Sternal Glands of found that aside from the last character, largely was consistent with his generic Scutelleridae (a-c) Coleotichus costatus (a) abdominal venter, sternal glands on SIV-SVI there was convergence in these character placements within tribal groups. LESTON (b-c) higher magnification of sternal glands states with other pentatomoid groups. (1952d, 1953a, latter reference with less hi- (d-f) Morbora schoutedeni (d) androconial erarchical arrangement) re-appraised the in- glands on SIV-SVI (e) higher magnification of microsculpture medial to androconial Infra-familial frafamilial classification of the scutellerids, glands (f) higher magnification of apex classification of the Scutelleridae with particular emphasis on the male aedea- cuticular duct of glandular unit of There are alternative views on the gus, introducing terminology for the ejacu- androconial glands (g-i) Tectocoris suprageneric classification of the Scutelleri- latory reservoir and conjunctival ap- diophthalmus (g) androconial glands on SIV-SVI (h) higher magnification of dae (Table 2). As with many other family- pendages. He maintained the suprageneric androconial glands on abdominal SVI (i) group taxa of the Heteroptera, their classifi- groups of Stål, but relegated a number of higher magnification of apex of cuticular cation derives from STÅL (1872, 1873), them as subtribes, within three scutellerid ducts of glandular units of androconial glands. Abbreviations: Ag = Androconial based on characters, such as, body contour, tribes (Eurygastrini, Pachycorini and Scutel- glands; Cd = cuticular ducts; Sg = sternal segment lengths of the labium, carination of lerini). In his conception, the male aedeagus glands. Scale bars = 0.1 mm. thoracic sterna, metathoracic glands, and of the Eurygastrini and Pachycorini are pregenital abdominal characters (e.g., more like those of the Pentatomidae, with stridulatory vittae, curvature of sternal su- the vesica bounded, sleeve-like, by the con- tures). He recognised seven suprageneric junctiva. He also recognised significant groups (suffix -aria), introducing three new within-group variation for the Pachycorini, groups (Elvisuraria, Eurygastraria and Odon- suggesting subgroups for Deroplax MAYR and totarsaria). SCHOUTEDEN (1904) mono- AMYOT & SERVILLE genitalic types. graphed the scutellerids, recognising the lat- He concluded that the Scutellerini sensu ter two suprageneric groups, and the Odon- Leston possess a derived male genitalic type, toscelaria as synonyms. KIRKALDY (1909) in characterised by: i. a cylindrical, sclerotized his world catalogue of the scutellerids, fol- phallotheca; generalized phallobase; sclero-

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tized vesica, with an apical positioned sec- other sternal glands that are putatively ho- ondary gonopore; two pairs of dorsal con- mologous in the elvisurine genus Coleotichus junctival appendages (CAI mostly membra- WHITE and Solenosthedium SPINOLA, and the neous with apical lobal sclerites, and CAII type genus of the Sphaerocoraria. Despite various, but mostly sclerotized) and a ven- these findings, CARAYON (1984) maintained tral pair of conjunctival appendages (CAI- the classification of LESTON (1952d), but el- II). MCDONALD (1961, 1963a, 1963b) in an evating the jewel bugs to family (as Scutel- investigation of the male and female geni- leridae), with all subordinate suprageneric talia of Australian jewel bugs, recognised taxa raised accordingly. SCHUH & SLATER the group as a family, and largely was in ac- (1995) adopted a modification of LESTON’s cord with LESTON. He suggested that a nov- (1952d) classification, recognizing four sub- el subtribal arrangement for Tectocoris families (Eurygastrinae, Odontotarsinae, HAHN might be required, on the basis of the Pachycorinae and Scutellerinae), but only genitalia of both sexes. KUMAR (1964, 1965) recognising a tribal classification within the disputed the findings of LESTON (1952d), nominotypical tribe. They provided a key concluding that the Eurygastrinae and and short diagnosis for each subfamily, plac- Pachycorinae do not have pentatomid male ing emphasis on the contour of the abdomi- genitalia, instead possessing an eversible nal sterna and hind wing venation. CASSIS ‘conducting canal’. Despite these re-inter- & GROSS (2002) adopted the system of MC- pretations and new observations, KUMAR re- DONALD & CASSIS (1984), recognising four tained the classification of LESTON (1952d), subfamilies for Australia, as well as outlining and in particular the subtribal arrangement the Eurygastrinae, Pachycorinae and Sphae- of Leston’s Scutellerini, but significantly, rocorinae as extralimital subfamilies, such moved the Australian genus Tectocoris, to that the Eurygastrinae are not divided into the largely Neotropical subfamily, the subordinate tribal groups (i.e. Eurygastrini, Pachycorinae. MCDONALD (1966) in a Odontoscelini and Odontotarsini). broad survey of genitalia of North American Pentatomoidea, illustrated many species, in The above-mentioned suprageneric more detail than previous workers for any groups of Scutelleridae have not been tested scutellerid taxon, and proposed equal rank- by phylogenetic analysis, with explicit char- ing for four of the suprageneric groups; acter definition and optimization, in the pri- namely, the Eurygastrinae, Odontotarsinae, mary literature. FISCHER (2001) in a pub- Pachycorinae and the Scutellerinae, with lished thesis gave character support for the no reference to the extralimital Elvisuraria monophyly of the Pachycorinae, Sphaero- and Sphaerocoraria sensu Leston. GROSS corinae, Elvisurinae, and a sister-group rela- (1975) subsequently recognised the elvi- tionship between Tectocoris and Odonto- surines as a subfamily, thereby recognising tarsinae. He also found that the Scutelleri- all the Australian jewel bugs within three nae and Odontotarsinae are not mono- subfamilies, the Elvisurinae, Odontotarsinae phyletic. It is beyond the scope of this work and Scutellerinae. MCDONALD & CASSIS to provide an analytical appraisal of these (1984) in a revision of the Australian groups, and discussions given below are scutellerids, confirmed these arrangements, couched within a narrative of character sys- but erected a new subfamily for Tectocoris tems, pending a cladistic analysis. Thus, we (Tectocorinae), on the aforementioned gen- have maintained the classification used in italic characters, as well as the presence of MCDONALD & CASSIS (1984) and CASSIS & male abdominal sternal glands. CARAYON GROSS (2002), and treatments below are (1984) provided a detailed anatomical ac- arranged alphabetically, as follows: Elvisuri- count of these latter glands, referring to nae, Odontotarsinae, Pachycorinae, Scutel- them as androconial glands. He found that lerinae and Tectocorinae. these glands were more widespread in scutellerids, with homologous structures also Taxonomic characters found in the Eurygastrinae sensu LESTON In this work we re-examined existing (Irochrotus AMYOT & SERVILLE, homologies and terminology used in the LAPORTE and Psacasta SPINOLA), as well as Scutelleridae. The characters examined in

284 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at this work use, in part, the homologies and species of Coleotichus. The eastern Aus- terminology established in MCDONALD & tralian species, C. artensis VAN DUZEE CASSIS (1984) and references therein, un- ranges from pale to dark brown, with the lat- less otherwise stated in discussions below. ter colouration type being influenced by the We have focused particularly on the exter- embrownment of the punctures. Species of nal efferent system of the metathoracic the extralimital species of Solenosthedium glands, male abdominal sternal glands, and exhibit more elaborate colouration, with the male genitalia. A survey of Australian bright yellow patterned markings against a and extralimital taxa was undertaken to pro- dark ground colour. vide a broader context for these decisions. In the Pachycorinae, many of the Colouration. The colouration of jewel species are dull in colouration, or can ex- bugs is generally diagnostic for species, al- hibit homogenous ground colour without though within some species there is extreme markings, such as in the genus FABRI- variation in colour and colour patterns. For CIUS. There are also spectacularly coloured example, in Tectocoris diophthalmus speci- species, such as the introduced species mens can vary from mostly orange to mostly Agonosoma trilineatum (Figs 16a, b), with metallic blue-green, which is further con- striped and spotted morphs. The Odonto- founded by partial sexual skewing, with tarsinae are mostly dull in colouration, in- males varying from mostly orange to mostly cluding the Australian species of Morbora, dark iridescent blue, whereas females are which is consistent with their epigaeic and notably paler, although ‘dominant-blue’ cryptozoic habits. morphs are not unknown (Figs 1f, 45a, b). In this work, we have documented The scutellerines are the most colourful of colour patterns of the pregenital abdominal all the jewel bugs, and the dorsum of these venter, which are often reliable for diagnos- species nearly always have a metallic-like ing species (Figs 21, 32). Although there is colouration, with polished and iridescent some intraspecific variation in these pat- orange, red, blue, green and purple hues, terns, we have found reliable discontinuities and most often with contrasting markings between related species. For example, (Figs 19, 24, 28). LYAL (1979) commented species of Choerocoris, which can be contin- on colour variation in Calliphara, proposing uous in the variation of the dorsum, have di- that light interference and pigmentation agnostic colour patterns of the venter (Fig. contribute to observed colouration. He re- 32). In Choerocoris grossi nov.sp., the venter ported that variation within a population of has a band of sublateral fuscous markings Calliphara species can be extreme, ranging (Figs 32a, b), which are absent in C. varie- from orange to blue/green individuals, and gatus (Figs 32g, h), although southwest that there are also geographical subspecies Western Australian populations of the latter of distinct colouration in species such as C. species sometimes have the segmental caesar (VOLLENHOVEN) and C. dimidiata boundaries weakly embrowned. (DALLAS). We have also observed consider- able variation in scutellerine taxa, such as In this work we do not treat the larvae, species of Lampromicra STÅL (Figs 28c-e) but note that the Elvisurinae, which gener- and Choerocoris DALLAS (Figs 24c-f). In par- ally have dull coloured adults, have apose- ticular, Lampromicra senator (FABRICIUS), matically coloured larvae in the genus varies from mostly iridescent green to purple Coleotichus, as in the Scutellerinae. or blue, and with or without an orange Texture. The texture of scutellerids is scutellar callus, and sometimes with darker highly variable; from smooth to highly patterning posteriorly. We urge caution in punctate, and less commonly rugo-punctate. using colouration in this species, as the pre- Many scutellerids are punctate, although in served colouration of museum specimens some cases this is deceptive as the punctures can be impacted by the ‘killing agent’ used. are often shallow. The dorsum can be more Species of Elvisurines are mostly pale to densely punctate on the underside of the dark brown (Fig. 3), with minor markings; body, and the distribution of punctures on e.g., minor black spotting in the Australian the thoracic pleura can be irregularly dis-

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tributed, In general, the Elvisurinae are lationships, aside from Calliscyta stalii, where punctate, but vary in the degree of the punc- the ventral surface of the tibiae has dense tation. For example, Austrotichus rugosus is and thick, yellow setae. The genital opening deeply punctate (Fig. 3a), with the punc- of the male pygophore is usually densely se- tures of the dorsum sometimes coalesced, tose, particularly on the ventral surface, and lateral regions of the pronotum and such as in Tectocoris diophthalmus (Fig. 46e, scutellum rugo-punctate. The other Aus- f). LYAL (1979) made significant use of the tralian species of elvisurines are all punc- setose patches on the genital opening in dis- tate, but often the punctures are shallower cerning species-groups within Calliphara. (Figs 3b-e). Most of the species of scuteller- We have not assessed the value of this char- ines are punctate, and deeply so in Choero- acter system across the Australian scutel- coris (Figs 24c-f) and Heissiphara minuta lerids, finding it in the first instance to be of nov.sp. (Figs 24b, 37). limited value. Whether these setose patches Vestiture. Few Australian scutellerids have a glandular function has not been de- have vestiture of taxonomic importance. termined. Extralimital exceptions include the species Head. The head of scutellerids has char- of the remarkable Afrotropical genus Elvi- acters of taxonomic and possibly phyloge- sura SPINOLA where the underside of the netic significance. In Agonosoma trilineatum, body is completely encased in scale-like se- the head is enlarged, being subequal to the tae. The vestiture of the Australian genus length of the pronotum (Figs 16a, b). Most Morbora DISTANT is diagnostic for the genus scutellerines have a triangular to subtrian- and the three known species (Fig. 13), and gular head, with the apices of the jugae ei- is comparable to the elaborate vestiture of ther just short or subequal to the anterior the extralimital odontotarsine genus Iro- edge of the clypeus. In many taxa the later- chrotus. Morbora have curly white sericeous al margins of the jugae are excavate; e.g., setae (Figs 13a, e, h) on the head, prono- particularly in Calliphara (Fig. 20a), and less tum, and thoracic pleura, which are inter- so in Choerocoris (Fig. 31a), Lampromicra mixed on the dorsum with fan-like (Figs (Fig. 39a) and Scutiphora (Fig. 42a). In con- 13c, h, i) or clove-like (Figs 13a, b, f) setae. trast, Cantao parentum has the margins rela- Most Australian species of Scutelleri- tively straight (Fig. 29a), although in ex- nae have the dorsum almost glabrous. The tralimital species, such as C. ocellatus notable exception is Lampromicra, which is (THUNBERG) the margins are a little exca- in part defined by a setose dorsum, and par- vate. The jugal margins may also be carinate ticularly so in L. senator (Fig. 1d). In the in lateral view, such as in species of Choero- Australian species, Cantao parentum, the coris (Figs 31a, b). Odontotarsines often head and pronotum are setose, and diagnos- have a more rounded head, including tic for this taxon. In the scutellerines, dense Morbora, which also has the lateral margins vestiture is also found in a number of ex- highly denticulate (Figs 11, 12a), although tralimital genera such as Brachaulax STÅL, it is not diagnostic at the species level. Procilia STÅL, Scutellera LAMARCK, and Antennae. The antennae of scutellerids Tetratharia DALLAS. Heissiphara minuta are relatively short, and articulated near or nov.sp. has setigerous punctures on both below the ventral margin of the eyes. ZRAZVY´ surfaces of the body. (1990) documented the evolution of anten- The antennae of most species jewel bugs nal structure in the Heteroptera, recognizing have simple, short setae, with the more the subdivision of the second antennal seg- proximal segments almost naked. The leg ment (= ) into pedicellites in the vestiture is also simple, with the femora Pentatomoidea. This is the case in the having scattered setae, mostly on the ven- Scutelleridae and we have introduced the tral surface. In nearly all scutellerids, the terminology of AII(a) and AII(b), for the ventral surface of the tibiae is more densely proximal and distal pedicellites respectively. setose, particularly more distally, as are the The shape of the antennae is often taxo- tarsi. In general the vestiture of antennae nomically diagnostic; e.g., AII(a) is arcuate and legs is not diagnostic or indicative of re- in Scutiphora pedicellata (Fig. 28b), particu-

286 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at larly in males. In most scutellerids AIII-AIV scutellerids have the anterolateral margins (occasionally AII as well) are flattened. The rounded in profile. The anterolateral mar- relative lengths of the antennae can be diag- gins vary in terms of their divergence, and nostic at both the generic and species levels. can be very strongly divergent, such as in For example, AII(a) or AII(b) are the short- Tectocoris diophthalmus (Figs 45a, b, 46a) to est segments in Calliphara (Figs 19a, b) and weakly divergent, as in Calliscyta stalii (Figs Scutiphora (Fig. 28b) respectively. 24a, 25a). In addition, the anterolateral margins are sometimes weakly excavate Labium. The labium is informative in (e.g., Tectocoris diophthalmus), to strongly terms of its absolute length, and relative excavate in extralimital genera of Odonto- proportions of the four segments. The labi- tarsinae (e.g., Odontotarsus LAPORTE). The um usually terminates between the apices of humeral angles are usually rounded, al- the mesocoxae (Fig. 4c) and abdominal SIV. though they are spinose in a few extralimital In those species with an elongate labium, species (e.g., ). The postero- the abdominal sterna are sometimes sulcate lateral margins are short and rounded. The serving as a housing for the labium; e.g., posterior margin is usually rectilinear in Elvisurinae species (particularly species of scutellerines, excavate in Tectocoris dioph- the extralimital genus Elvisura SPINOLA) and thalmus (Figs 47a, b), to strongly rounded in Tectocoris diophthalmus. STÅL (1873) and the elvisurine genus, Coleotichus (Figs 3c-e). SCHOUTEDEN (1904) afforded value to the relative lengths of the labial specimens, par- Scutellum. All scutellerids have a great- ticularly LII in relation to subsequent seg- ly enlarged scutellum, which reaches near ments, in discerning suprageneric groups. the lateral margins of the abdomen, with We find that there is significant variability the areas proximal to the forewing articula- in such characters, even within genera (e.g., tion and abdominal connexiva exposed Coleotichus). (e.g., Figs 3, 16, 19, 24, 28, 37 and 45a, b). The scutellum is strongly convex in profile, Pronotum. The pronotum is relatively and in some species is very strongly de- uniform in the scutellerines, and is mostly clivent posteriorly, such as in Tectocoris dio- trapezoidal in shape (e.g., Figs 3, 16, 19, 24, phthalmus (Fig. 47). The anterior margin of- 28, 37 and 45a, b). Some genera of the ten has sublateral foveae, however it is not Odontotarsinae, including the Australian regarded as having classificatory signifi- genus Morbora, and extralimital species of cance; occurring in unrelated taxa such as Irochrotus, are exceptional in the scutel- Austrotichus rugosus and Cantao parentum lerids, where the lateral margins are not (cf. Figs 3a and 28a). The extent of scutellar clearly differentiated into anterior and pos- coverage of the hemelytra and abdomen has terior components, being more rounded taxonomic significance. In many scuteller- across the entire length, with the anterior ines, the hemelytral membrane extends be- angles projected in front of the posterior yond the scutellum at rest (e.g., Figs 19a-d margins of the eyes (e.g., Morbora: Figs 11, and 28a), with a few exceptions including 12b, extralimital Irochrotus). The contour of the species of Choerocoris, Calliscyta stalii the pronotum is usually strongly convex, and Heissiphara minuta nov.sp. (Figs 24a-f). and declivent towards the posterior margin. In the extralimital genus Eurygaster LA- The pronotum is often divided into a weak- PORTE, the scutellum is diagnostically less ly demarcated callosite region, with sublat- extensive, with the forewing more exposed. eral muscle attachment ‘scars’. In a few taxa, such as Lampromicra senator, the callosite re- Thoracic Sterna. Characters of the tho- gion is demarcated by a broad transverse fur- racic sterna are most important in differen- row (Fig. 28e). The anterior margin varies tiating the Elvisurinae from the remainder between weakly (e.g., Figs 28a, 29a) to of the Scutelleridae. Elvisurines have the strongly concave (e.g., Fig. 9a) conditions. thoracic sterna greatly modified into keel- The anterolateral margins are sometimes di- like structures, which are relatively thin and agnostic, and can be explanate, with the elevated in species of Coleotichus ( Fig. 7c) margin carinate (Figs 25c, 46b), and if so, to thick and moderately high, as in Aus- the head is usually also carinate. Some trotichus rugosus (Fig. 4c) and Solenotichus

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circuliferus (Fig. 9c). In Coleotichus, the Within the Pentatomoidea, the external mesosternal keel is anteriorly projected and efferent system has been under-utilized in overlaps the prosternal keel. In other scutel- defining supraspecific groups. This character lerids the thoracic sterna are flat to sulcate, system was used in the suprageneric classifi- and exhibit little variation. The most no- cation of scutellerids, as far back as STÅL table exceptions exist in the Eurygastrinae (1873), who differentiated his notion’s of and Odontotarsinae, where the proepister- the Eurygastraria and Odontotarsaria, on nal anterior edge can be explanate and con- the basis of distinct and indistinct ‘orifices’ tiguous with the prosternum, but never of respectively. SCHOUTEDEN (1904) also the height found in elvisurines. In some recognised that the Pachycorinae (as odontotarsines, the mesosternal margins can Tetyraria) had weakly developed orifices. also be a little raised, housing the labium, We have found that variation in the per- but not prominently as in elvisurines. itreme is taxonomically significant; poorly developed to almost absent in most species Thoracic Pleura. The thoracic pleura of Odontotarsinae and Pachycorinae (ex- are relatively uniform, aside from the anteri- ceptions include Morbora and Agonosoma in or margin of the proepisternum, the cover- each of these subfamilies, where the per- age of the evaporative areas, and the condi- itreme is prominent). Other examples in- tion of the external efferent system of the clude the extralimital Sphaerocorinae (both metathoracic glands. In scutellerids the an- Hyperoncus STÅL and Sphaerocoris BURMEIS- terior margin of the proepisternum is nearly TER), where the peritreme is greatly elongate always prominent and explanate, which in and narrow, subparallel to the posterior mar- the elvisurines is contiguous with the gin of the mesepimeron. prosternal keel (e.g., Fig. 7c). In most taxa, this margin is explanate, housing the eye, The condition of the peritreme and ex- except in Tectocoris diophthalmus. The pres- tent of the evaporative areas are most in- ence or absence of evaporative areas of the formative at the genus level. For example in posterior margin of the mesepimeron elvisurines, Coleotichus possesses a linear, (bounding the metathoracic spiracle) is also elongate peritreme, which is sulcate medial- informative at a taxonomic level, such as in ly (Fig. 7d). Solenotichus also has an elongate Calliscyta stalii where such areas are absent. peritreme, but is strongly raised (almost au- The absence of evaporative areas on the riculate) distally. In comparison, Aus- mesepimeron appears to be correlated with trotichus has a relatively short peritreme reduction in the external efferent system of (Fig. 4d), which is also true of the extralim- the metathoracic glands. ital genus Solenosthedium, and in Elvisura, the peritreme is near obsolete. Many of the Metathoracic Scent Glands. The exter- scutellerines have a subreniform-shaped nal efferent system of the metathoracic peritreme, which can be large (e.g., Cal- glands is informative at a diagnostic and liphara, Figs 20c, d); Lampromicra, Fig. 39d) phylogenetic level in the Heteroptera (e.g., to small (e.g., Scutiphora, Fig. 42d). In con- CARAYON 1971: Heteroptera; KELTON 1978: trast, other genera have an obovate per- , SLATER 1979: , CAS- itreme, which is more expanded laterally, SIS 1995: Miridae). The terminology used such as Choerocoris spp. (Fig. 31d) and Heis- for the pterothoracic pleura, particularly for siphara (Fig. 38c). In Calliscyta STÅL (Fig. characters of the metathoracic gland effer- 25d) and Cantao AMYOT & SERVILLE (Fig. ent system follow those given for the Miri- 29d), the peritreme is greatly enlarged, sub- dae (Heteroptera) in CASSIS (1995). In this triangular and polished, occupying more work, the following definitions are applied: than 1/2 of the metepisternum, and the i. ostiole: opening of the metathoracic evaporative areas are reduced in extent. In glands; ii. peritreme: the canal emanating Tectocoris, the external efferent system is from the ostiole; and, iii. evaporative areas: greatly reduced to a linear, sulcate strip, ‘mushroom-like’ bodies bounding the per- with few evaporative bodies (Fig. 46d). itreme and ostiole on the metepisternum. The position, shape, size, and orientation of Pregenital Abdomen. The morphology the ostiole, peritreme and evaporative areas of the pregenital abdomen is most informa- can all be informative. tive at the generic and species levels. The

288 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at pregenital abdomen has five important char- rative areas or evaporatoria) (CARAYON acters that we examined: sternal glands (Figs 1971; COBBEN 1978; STADDON 1979; 2a-i) and stridulatory structures (Figs 17e-f), ALDRICH 1988; SCHUH & SLATER 1995). which are informative at the suprageneric More recently the discovery of various level; the presence or absence of posterolat- unicellular abdominal sternal glands has eral processes, which are informative at both yielded morphological and anatomical evi- the generic and specific levels; the exposure dence that has taxonomic significance, in of the eighth urite; and, the development of disparate groups such as the : SVII, which can cover both the male and fe- Holoptilinae (WEIRAUCH & CASSIS in male terminalia in Agonosoma trilineatum. press), and numerous families and infrafa- In the Pachycorinae, both sexes have milial groups of the Pentatomoidea (CARAY- stridulatory structures found sublaterally on ON 1981; STADDON & EDMUNDS 1991; abdominal SV-VI (plectrum on metatibiae), STADDON 1999; FISCHER 2000). In the Pen- as seen in Agonosoma trilineatum (Figs 17e- tatomoidea, many of these glandular systems f). In the extralimital genus Vanduzeenia have been found in males, and have been SCHOUTEDEN, originally placed in the determined to be concerned with the pro- Odontotarsinae, we found stridulatory areas, duction of sex (STADDON et al. medially positioned on SIII-SVI, consoli- 1987; STADDON 1990; ALDRICH 1996). dating its place in the Pachycorinae. No CARAYON (1984) described a novel sternal other scutellerid of the other subfamilies gland system, which he termed androconial known to us possess stridulatory areas on the glands, in the males of scutellerids; in detail abdominal sterna. for the Australian species, Tectocoris dioph- thalmus, and in some extralimital odontotar- The lateral margins of the pregenital ab- sine species of Irochrotus, Odontoscelis and domen possess spines of nodules that are di- Psacasta; all of which he regarded as homol- agnostic. In the genus Calliphara, species ogous. He also reported androconial glands vary in the possession of spines on the pos- in the elvisurine taxa, Solenosthedium biluna- terolateral angles of the pregenital abdomi- tum (FABRICIUS) and Coleotichus costatus nal sterna, varying in combination from (FABRICIUS) (as C. unicolor (FABRICIUS)). SIV-SVII (Figs 21a, b), and are sometimes These two types are putatively homologous, absent (e.g., Fig. 21c). The connexiva are but we regard them as distinct enough, to also diagnostic for species of Choerocoris; C. warrant different terminology, which we lattini nov.sp., has the posterior angles nodu- propose as follows: i. androconial glands late (Fig. 32c, d), unlike the other Choero- (with peg-like ducts; e.g., Figs 2d-f, g-i), coris species (Figs 32a, b, e, f). which are found in the Odontotarsinae and In most cases the eight abdominal urite Tectocorinae; and, ii. setose sternal glands is recessed and not visible. In the unrelated (with elongate setae; Figs 2a-c), which are species Cantao parentum and Tectocoris dio- found in the Elvisurinae. We also found that phthalmus, SVIII is exposed, covering the all the species of the Australian odontotar- posterior margin of the male pygophore sine genus Morbora have androconial glands (Figs 29e, 46e). (Figs 2d-f), that are remarkably alike in ex- ternal structure and distribution to those Abdominal Sternal and Androconial found in Tectocoris diophthalmus. Glands. The presence of multicellular dor- sal abdominal glands in larvae and most STADDON (1999) reported the presence adults of the Heteroptera are defining for of male sternal glands in two British species the suborder, and have significant classifica- of Eurygaster, with two types of unicellular tory value within the suborder (DUPUIS glands, one of which he postulated as possi- 1947; ALDRICH 1988; SCHUH & SLATER bly being androconial glands. Under light 1995). The multicellular metathoracic microscopy, we were able to discern mi- glands are diagnostic for the Neo- crotrichiae broadly distributed across the heteroptera, but particularly in the Lep- abdominal sterna of these two species, but topodomorpha + Trichophora, which have no structures that appeared like androconial the characteristic mushroom bodies (evapo- or setose sternal glands.

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Table 3: Presence, extent and type of male androconial glands in Scutelleridae. ssg = Morbora (SIV-SVI), and Irochrotus and setose sternal glands, ag = androconial glands. + = fused glandular patches; ++ = paired Odontoscelis (SV-SVI). In the Tectocorinae glandular patches. the androconial glands are found on abdom- Taxon Gland type SIII SIV SV SVI SVII inal SIV-SVI. The setose sternal glands of Elvisurinae the Elvisurinae are found in all species of Coleotichus biroi ssg ++ ++ ++ Solenosthedium examined, but vary in their Coleotichus costatus ssg ++ ++ ++ distribution, from SIII-SVII to SIV-SVII, Coleotichus fuscus ssg ++ ++ ++ and in all cases are densely setose, more so Solenosthedium chinense ssg ++ ++ ++ ++ than any other elvisurine examined. The se- Solenosthedium lederei ssg ++ ++ ++ ++ ++ tose sternal glands are found in some species Solenosthedium liligerum ssg ++ ++ ++ ++ ++ of Coleotichus, but are missing in many Solenosthedium lyriceum ssg ++ ++ ++ ++ ++ species examined (C. artensis, C. blackburni- Solenosthedium madagascarense ssg ++ ++ ++ ++ ae WHITE, C. bulowi SCHOUTEDEN and C. Solenosthedium rubropunctatum ssg ++ ++ ++ excellens WALKER); in most species where Solenosthedium schulzi ssg ++ ++ ++ ++ ++ these glands are present, they are distributed Solenotichus circuliferus ssg ++ ++ ++ + on SIV-SVI, where the setae are moderately Odontotarsinae dense. Setose sternal glands are present in Irochrotus incisus ag ++ ++ the monotypic genus Solenotichus MARTIN, Irochrotus indicus ag ++ ++ where they are medially contiguous on Irochrotus mongolicus ag ++ ++ SVII. We found no evidence of these glands Morbora australis ag ++ ++ ++ in Austrotichus and Elvisura. Morbora hirtula ag ++ ++ ++ Morbora schoutedeni ag ++ ++ ++ The phylogenetic value of setose sternal Odontoscelis byrrhus ag ++ ++ and androconial glands requires a broader Odontoscelis dorsalis ag ++ ++ sampling within the Scutelleridae. FISCHER Odontoscelis fuliginosa ag ++ ++ (2001) found the androconial glands to be Odontoscelis iberica ag ++ ++ synapomorphic for Tectocoris + Odontotarsi- Odontoscelis litura ag ++ ++ nae. The of the setose and andro- Odontoscelis lineola ag ++ ++ conial glands needs to be investigated fur- Odontoscelis signatus ag ++ ++ ther, so that the anatomical features of the Periphima batesoni ag ++ ++ ++ ++ ++ glandular units are described. Our current Psacasta cerinthe ag ++ ++ ++ ++ + hypothesis is limited by observation of only Psacasta exanthematica ag ++ ++ ++ ++ ++ the external efferent system component of Psacasta pallida ag ++ ++ ++ ++ ++ the elvisurine setose sternal glands. In addi- Tectocorinae Tectocoris diophthalmus ag ++ ++ ++ tion, we need a broader understanding of the distribution of these gland types within In Table 3, we list scutellerid taxa that the scutellerids. we examined which possess sternal glands of Female Terminalia. The terminology of the two types described, and their distribu- female genitalia follows that given by MC- tion on the abdominal sterna. The results of DONALD (1966), and the homologies and this survey indicate that the androconial terminology therein have not been re- glands, are widespread in the Odontotarsi- assessed in this work. In the main, the ex- nae, but not universal; no cuticular evi- ternal plate-like sclerites do not differ signif- dence of such glands were found in Alpho- icantly from other pentatomoids, with ellip- coris GERMAR, Ceatocranum JAKOVLEV, Ellip- toid to triangular paratergites VIII and IX, socoris MAYR, GERMAR, Polyphy- and subtriangular gonocoxae I. In scutel- ma JAKOVLEV, Promecocoris JAKOVLEV and lerids there is variation in the orientation of Xerobia STÅL. Androconial glands were these sclerites, which are either ventral in found in the species of Irochrotus, Morbora, orientation, such as in elvisurines and Odontoscelis, Periphima JAKOVLEV and scutellerines such as Calliphara and Lam- Psacasta that we observed. We found varia- promicra, to caudal in orientation, as in Cho- tion in the distribution of the androconial erocoris and Morbora. The orientation of glands, from widespread in Psacasta and Pe- these sclerites can also vary relative to each riphima (SIII-SVII), to less widespread in other; they can be co-planar and flat such as

290 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at in Coleotichus to bi-planar, where parater- mus has a unique spermatheca, with the gites are obtusely oriented relative to gono- pump minute, and the bulb with an arcuate, coxae I. In Choerocoris species the size and acute apex. shape of the female terminalia sclerites are Pygophore. The pygophore has a num- of great importance in separating species. ber of characters that have diagnostic signif- Spermatheca. We re-examined the sper- icance. The pygophore varies in orientation, matheca of the Australian species, and found and can be ventral (e.g., elvisurines, Cal- no significant enhancement of morphologi- liphara spp., Tectocoris diophthalmus; Figs 4e, cal understanding beyond MCDONALD & 20e, 46e) or caudal in orientation (Choero- CASSIS (1984); therefore, no illustrations of coris spp., Figs 32b, d, f, h). LESTON (1952a, the spermatheca are provided in this work 1952b), regarded the presence of a row of and users are referred to the latter reference. peg-like bristles on a pygophoral process In most scutellerids, the spermatheca has a defining for the Sphaerocorinae. This struc- membraneous to lightly sclerotized fecunda- ture has not been found in any of the Aus- tion canal, a dilated reservoir, and a pump tralian scutellerids. In addition, LYAL (1979) composed of flanges to which muscles at- considered the distribution of setae on the genital opening of the male pygophore to be tach, and a distal bulb. FISCHER (2001) pro- of importance in determining species-group poses an alternative nomenclature, rejecting boundaries. We found that the distribution the usage of ‘spermathecal reservoir’, on the of setae on the pygophore was not particu- basis that there is no evidence of a ‘reservoir’ larly noteworthy for the Australian taxa, function for this character state. He uses the and did not use it in our classification. terminology of the ‘dilation of the spermath- ecal duct’. Similarly, he rejects the function- Parameres. The parameres, as with all al description of a ‘spermathecal pump’. We other pentatomoids, are symmetrical, and did not examine these character systems in small to relatively large. The stem of the any great detail, and prefer to use the de- parameres is nearly always columnar, and scriptive terms used in MCDONALD & CAS- thin to thick. In some cases, such as in some SIS (1984), recognising their limitation. elvisurines (e.g., Austrotichus rugosus, Coleo- tichus species; Figs 5b, 7b) and the scuteller- The base of the spermatheca has scle- ine genus Choerocoris (Figs 35b, 36b) the rites bounding the opening of the fecunda- apex of the stem has a lobe-like flange. tion canal of the spermatheca. GAPUD Nearly all species have a hook-shaped (1991) regards this as diagnostic for scutel- crown, which can be greatly enlarged, as in lerids, a view supported by Grazia (pers. Tectocoris diophthalmus (Fig. 47b). There ap- comm.). There is variation in these scle- pears to be no phylogenetic or suprageneric rites, as reported in MCDONALD & CASSIS classificatory value in the condition of the (1984), but have not been further investi- parameres, although they are sometimes di- gated in this work. The fecundation canal agnostic at the species level. can be short (e.g., Choerocoris spp.) to elon- Aedeagus. Terminology and homologies gate (e.g., Coleotichus costatus), although its of the male aedeagus of pentatomoids are length is not necessarily informative above various (e.g., SINGH-PRUTHI 1925; LESTON the species level (e.g., Coleotichus artensis). 1952d; DUPUIS 1955; KUMAR 1964, 1965; The spermathecal reservoir is most often MCDONALD 1966; AGARWAL & BAIJAL round in scutellerines, and relatively large, 1984; TSAI et al. 2004), and we use the fol- with a few exceptions where it is narrow lowing terminology, including some modifi- (e.g., Austrotichus rugosus GROSS, Morbora cations of the above authors, as follows: spp.). The spermathecal reservoir often has a ribbed substructure, as in most scuteller- i. phallobase (= articulatory apparatus) ines, but can be simple and uniformly mem- which is a U-shaped process attached to the mesial surface of the ventral aspect of the braneous (e.g., Austrotichus rugosus). The pygophore, with generalised capitate pro- bulb is also uniform in the Australian scutel- cesses, and housing the primary gonopore. lerids, round and unmodified. However, in Morbora species the bulb is more elongate, ii. ductus seminis proximalis, is usually a with its margins sinuate. Tectocoris diophthal- narrow, membraneous tubule, originating

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from the primary gonopore. In a few taxa, ing canal (= ‘lumen ‘of TSAI et al. 2004), the ductus seminis proximalis is broader, which is largely undifferentiated, continuous with light sclerotization, and bounded by a with the ductus seminis distalis, and without membraneous tubule, as in Agonosoma trilin- a discrete separation of the two. For utilitar- eatum (Figs 18c, d) and Tectocoris diophthal- ian purposes and realising its limitations, we mus (Figs 47c, d). Tsai (pers. comm.) regards refer to the commencement of the ductus the outer tubule as derived from a ligament seminis distalis, as distal to the junction of of the phallobase, and we introduce the the ventral conducting canal and the ductus term ‘ligamentary tubule’ for this structure. seminis proximalis. iii. the term ‘ejaculatory apparatus’ is iv. the vesica is a distal tubule bounding proposed for the medial modification of the the ductus seminis distalis, and is attached ductus seminis, and begins at the apical basally to the conjunctival appendages. opening of the ductus seminis proximalis, There is serious doubt that the notion of the and basad of the distal margin of the phal- vesica, introduced by SINGH-PRUTHI (1925), lotheca and vesica. This is synonymous with is homolgous to that found in the Pentato- the ejaculatory reservoir of SINGH-PRUTHI momorpha and the . KERZH- (1925) and subsequent workers (e.g., MC- NER & KONSTANTINOV (1999) have empha- DONALD 1966; TSAI et al. 2004), the ‘semi- sised that in the Miridae, the vesica is a dis- nal reservoir’ of BAKER (1931), and the ‘con- tal differentiation of the endosoma, and de- ducting chamber’ of KUMAR (1964). The fined by its positional attachment to the ejaculatory apparatus includes the dorsal and secondary gonopore, and functionally by its ventral conducting canals of Kumar, and the method of eversion or inflation. In contrast, sac-like intermediary chamber, which is of- KUMAR (1965) hypothesised that the vesica ten labeled as the ‘ejaculatory reservoir’. In was derived from the conjunctiva; this could Figures 5c & d these constituent parts are be interpreted as being in contrast to endo- first labeled, and then throughout the publi- somal differentiation. In the Scutelleridae, cation. The ventral conducting canal is we retain the use of the vesica, largely as a basally oriented, subparallel to the ventral descriptive term, and in the main is a scle- margin of the phallotheca, and usually scle- rotized tubule which parallels the contour of rotized. The canal is often convoluted, in an the ductus seminis distalis, although it can alternating pattern, giving the impression have subdistal or apical processes, as in some that the canal is paired, as interpreted by species of Choerocoris (e.g., Fig. 36c) and KUMAR (1964, 1965). MCDONALD (1966) Calliphara (e.g., Figs 22c, d) respectively. In has shown this canal to be undivided in the latter case, the secondary gonopore is cross-section. The number of convolutions subapical, but exhibits no further complexi- in the ventral conducting canal is sometimes ty. The condition of the vesica is very differ- diagnostic for taxa, such as in Cantao (e.g., ent in some pachycorinae species, including, Figs 30c, d) where there are as many 12 con- Agonosoma trilineatum, where it is fully volutions, whereas in some species of Cal- membraneous and broadly attached to the liphara there are no apparent convolutions phallotheca (Figs 18c, d). (e.g., C. regalis (FABRICIUS), Figs 22c, d). The convoluted ventral conducting canal is v. the conjunctival appendages are characteristic of species of the Elvisurinae processes that are attached mesially to the (e.g., Figs 5c, d), Scutellerinae and Sphaero- vesica, and externally to the phallotheca, corinae, but is not unknown in other sub- and are often fused basally to each other. LE- families, aside from the Tectocorinae (Figs STON (1952d) introduced a terminology for 47c, d). The ejaculatory reservoir varies in the conjunctival appendages, recognising shape, from oval to elliptoid in shape, and in three pairs in the Scutelleridae. He described size, from large (subequal in length to the two dorsal pairs of conjunctival appendages, ventral conducting canal) to relatively which are basally contiguous, and termed small, as well as being lightly to heavily scer- them as the first and second pairs; and also, lotized (Fig. 47c). The ejaculatory reservoir a ventral pair, the so-called third conjuncti- terminates adjacent to the ventral conduct- val appendages. We have modified this ing canal, and enters into a dorsal conduct- nomenclature, regarding the first and second

292 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at pairs as the second pair only (our CAII), (e.g., Figs 7c, 25b, 46c); antennae four-seg- which are most often subdistally bifurcate, mented, AII divided, distal segments often with lateral (CAII(L)) and medial branches flattened; labium four-segmented, most of- (CAII(M)). These are regarded as the sec- ten reaching between metacoxae and ab- ond conjunctival appendages, because in dominal SIV; pronotum trapeziform, with some cases, there is a pair of conjunctival ap- anterolateral and posterolateral margins pendages which are more ventrad. We refer (e.g., Figs 3, 16, 19, 45a, b), or rounded lat- to these as the first conjunctival appendages eral margins (Figs 16a-c); callosite region (CAI), and they can also be branched, as in weakly demarcated; scutellum enlarged, some Calliphara species, where there are dor- covering most of abdomen and wings, with sal (CAI(D)) and ventral (CAI(L)) branch- exocorium and clavus minimally exposed es (e.g., C. regalis, Figs 22 c, d), or they can proximally (e.g., Figs 3, 16, 19, 45a, b); be membraneous outgrowths of the phal- forewing membrane sometimes exposed be- lotheca as in Morbora schoutedeni DISTANT yond tip of abdomen (e.g., Figs 19a-d); frena (Figs 14c, d) and Tectocoris diophthalmus (Figs reduced or absent; hemelytral membrane 47c, d). The shape, size, and degree of scle- with 10 or more longitudinal veins; external rotization of each of the conjunctival ap- efferent system of metathoracic glands pres- pendages are diagnostic at the species, and ent (e.g., Figs 4d, 17c, 20d, 46d); ostiole sometimes generic levels. The CAII are of- placed anteriorly on metepisternum; evapo- ten useful in scutellerid classification, where rative areas present on metepisternum, and the lobal sclerites may be short and conical most often on mesepimeron (e.g., Figs 4d, (e.g., Figs 5c, d, 10c, d), arcuate (e.g., Figs 8c, 12d, 20d, 31d); parameres with a columnar d, 14c, d, 26c, d), or denticulate (e.g., Figs stem, and usually with a hook-shaped crown 40c, d). The CAIII are also various, and can (e.g., Figs 5b, 22b, 47b); aedeagus most of- be separated (e.g., Figs 36c, d), or partially to ten with an ejaculatory apparatus (e.g., Figs fully contiguous medially (cf. Figs 8c, d, 14c, 5c, d, 26c, d, 30c, d); first and second go- d, 26c, d). On the basis of the Australian napophyses membraneous; spermathecal scutellerids, we have found no features of the reservoir without a sclerotized rod; and, conjunctival appendages that support exist- spermathecal bulb round without processes. ing suprageneric groups. Description: Body elongate to elongate- ovoid; moderately to strongly biconvex, sometimes dorsum more rounded than ven- ter; mostly large species, between 8-20 mm, Family Scutelleridae smallest species < 4 mm, largest species > 20 mm; body either dull brown to black in Scutellerida LEACH 1815: 123 (suprageneric group) colouration (e.g., Figs 3a-e), or brightly Scutelleroides: DALLAS 1851: 3 (superfamily; in- coloured, with iridescent hue and often with frafamilial classification) contrasting markings (e.g., Figs 19a-d, 24a-f, Pentatomidae, Scutellerina: STÅL 1872: 32 (clas- 28a-e); body often punctate, with shallow sification); STÅL 1873: 3 (classification) (e.g., Figs 7a), to deep punctures (e.g., Figs Pentatomidae, Scutelleridae (sic): LETHIERRY & 4a, 12a, 31a, 37), usually regularly distrib- SEVERIN 1893: 15 (catalogue) uted, sometimes punctures coalesced (e.g., Pentatomidae, Scutellerinae: SCHOUTEDEN 1904: Fig. 3a); body less commonly rugo-punctate 1 (monograph); KIRKALDY 1909: 263 (catalogue); on lateral aspects of pronotum and abdomi- LESTON 1952d: 13 (subfamily) nal venter; body mostly with sparse distribu- Scutelleridae: MCDONALD 1966: 67, 68 (geni- tion of simple setae, rarely with dense distri- talia); MCDONALD & CASSIS 1984: 550 (revi- bution of simple setae (e.g., Fig. 1d), rarely sion); CASSIS & GROSS 2002: 580 (catalogue) with setigerous (Figs 31a, 38a) or scale-like Diagnosis: Scutellerids are recognised by setae (e.g., Figs 13c, h, i). Head: usually tri- the following combination of characters: angular to subtriangular (e.g., Figs 4a, 7a, body moderately to strongly biconvex (e.g., 17a) sometimes suboval (e.g., Fig. 9a); jugae Fig. 1); apices of jugae and anterior edge of large, not exceeding the anterior edge of clypeus subequal in length (e.g., Figs 4a, 7a, clypeus (e.g., Figs, 4a, 20a, 42a, 46a); lateral 20a, 46a); bucculae flattened, sub-parallel margins of jugae most often excavate (e.g.,

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Figs 20a, 25a), sometimes linear (e.g., Figs sometimes exposed, tip of hemelytral mem- 29a, 46a), rounded (e.g., Figs 7b, 25c, 38b, brane sometimes visible beyond scutellum at 39b, 42b) or carinate (e.g., Figs 29b, 31a, b) rest (e.g., Figs 19a-d, 28a); strongly convex, in profile; clypeus well-developed, lateral often strongly declivent posteriorly, beyond margins reaching between frons (e.g., Figs midpoint; anterior margin sometimes with 4a, 7a, 9a, 31a) and anterior aspect of vertex sublateral foveae (e.g., Figs 24a, 28a); later- (e.g., Figs 25a, 38a, 42a); vertex weakly con- al margins weakly to strongly rounded, vex; lorae triangular, not visible from above, sometimes strongly divergent posteriorly; tip with strongly demarcated margins; bucculae rounded, rarely truncate (Fig. 24a). Frena: large, narrow, subparallel, usually reaching greatly reduced to absent. Hemelytra: mem- near base of head (e.g., Figs 4c, 7c, 20c, 31c, brane with ten (e.g. Fig. 24c) or more longi- 46c); gula short, convex. Eyes: contiguous tudinal veins. Thoracic pleura: anterior with pronotum (e.g., Figs 3, 24, 45a, b); usu- margin of proepisternum weakly (e.g., Figs ally moderately sized, ≥ 1/3 of head length, 4c, 7c, 20c, 31c) to strongly (e.g., Figs 7c) sometimes large, about 1/2 head length (Fig. explanate, sometimes excavate at eye (e.g., 19d); pair of ocelli, removed from posterior Fig. 7c), sometimes entire and linear (Figs margin of head (e.g., Figs 3, 24, 45). Anten- 17b, 39c, 46b-c); mesepimeron subrectangu- nae: inserted anteriad of ventral margin of late, most often with evaporative areas on eyes (e.g., Figs 7b, 9b, 20b, 29b, 31b, 38b, posterior margin, bounding metathoracic 39b, 46b); four-segmented, AII divided, rel- spiracle (e.g., Figs 4d, 7d, 9d, 12d, 17c, 20d, atively short; segments linear, cylindrical, 31d, 38c, 42d), sometimes extending to AII(b)-AIV often flattened; AII rarely arcu- meso-supracoxal lobe, sometimes absent ate; AI short, sometimes AII(a) and AII(b) (e.g., Figs 25d, 46d), posterior margin some- shortest segment; AIV often longest seg- times swollen (e.g., Fig. 12d); metepister- ment. Labium: usually reaching between num usually with well-developed external apices of mesocoxae to abdominal SIV, efferent system of metathoracic glands (e.g., rarely longer; LI reaching posterior margin Figs 4d, 7d, 9d, 17d, ) usually covering at of head; LII usually longest segment; LII least 1/2 of segment, rarely reduced (e.g., sometimes laterally bicompressed; LIII-LIV Figs 12d, 42d, 46d); ostiole small (e.g., Fig. sometimes dorsoventrally flattened. Prono- 4d, 46d) to moderately large, oriented near tum: large, trapeziform; anterior margin anterior margin of segment; peritreme very weakly (e.g., Figs 3a, 7a) to strongly concave small (e.g., Fig. 4d) to greatly enlarged (e.g., (e.g., Fig. 9a); anterolateral margins elon- Fig. 7d), either linear (e.g., Figs 7d), linear gate, weakly (e.g., Figs 16a, 25b) to strongly with apex anteriorly recurved (e.g., Figs 17c, divergent (e.g., Figs 9a, 19a-d, 45a,b, 46a) 31d), subreniform (e.g., Figs 20d, 39d, 42d), posteriorly, anterolateral margins usually obovate (Fig. 38c) or greatly enlarged and linear (e.g., Figs 3a-e, 19a-d), rarely weakly subtriangular (Figs 25d, 29d), peritreme of- convex (e.g., Figs 24c, d, e) or concave (e.g., ten medially sulcate (e.g, Figs 7d); evapora- Figs 28b, 45a, b), carinate (e.g., Fig. 25d) or tive areas reduced (e.g., Figs 25d, 29d, 42d, rounded in profile; posterolateral margins 46d) to extensive (e.g, Figs 4d, 7d, 9d, 17c, short, most often weakly convex; posterior 31d), often extending laterally beyond tip of margin usually rectilinear (e.g., Figs 3a, 16a, peritreme. Thoracic sterna: usually flat (e.g., b, 19a-d, 24a-f), rarely rounded (e.g., Figs Figs 12c, 20c, 29c, 31c, 39c, 42c, 46c), 3c-e), if so covering anterior margin of sometimes lateral margins strongly raised, scutellum, rarely weakly excavate (e.g., Fig. keel-like (e.g., Figs 4c, 7c, 9c), rarely with 28a); callosite region with sublateral trian- pro- and mesosternal keels overlapping (Fig. gular markings, most often not strongly de- 7c). Legs: femora fusiform; tibiae terete, tar- marcated, co-planar with pronotal disc (e.g., si three-segmented. Pregenital Abdominal Figs 3a-e), rarely with transverse furrow Venter: SII-SVII always visible (Figs. 21a-d, (e.g., Figs 28c-e). Scutellum: greatly en- f-i), SVIII most often recessed, rarely ex- larged, shield-like, U-shaped, covering most posed in males (Figs 21e, 45c), covering pos- of hemelytra and abdomen (e.g., Figs 3a-e, terior margin of pygophore (Figs 29e, 46e); 16a, b, 19a-d, 24a-f, 28a-e, 45a, b), exocori- sometimes posterolateral angles with acumi- um always partly exposed, base of clavus nate spine (e.g., Figs 21a, b); males with

294 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at sternal glands sublaterally, ranging between 14c, d, 43c, d, 47c, d), sinuate (e.g., Fig. 5c, SIII-SVII (Figs 2a-i, 12e); sometimes males d, 8c, d, 30c, d, 35c, d, 40c, d), rarely sub- and females with stridulatory region on SV- distally strongly incrassate (e.g., Figs 30c, SVI (Figs 17e-f). Male Genitalia: pygophore 35c), sometimes with subdistal processes moderate size to large, ventral surface cau- (Fig, 36c, d), or with distal process above dally (e.g., Figs 4e, 9e, 12e, 31e) or ventral- secondary gonopore (Figs 22c, d); CAI most ly oriented (e.g., Figs 20e, 29e, 42e, 46e); often absent (e.g., Figs 5c, d, 8c, d, 10c, d, ventral margin convex, emarginate or sub- 26c, d, 30c, d, 35, d, 36c, d, 40c, d, 43, d), if linear; genital opening moderately broad to present medially fused at base (Fig. 14c, d, broad, with setose regions usually on dorsal 47c, d), or bifurcate with ventral and dorsal margin (e.g., Figs 22a, 43a), and often on branches (e.g., Figs 22c, d); CAII sometimes ventral (e.g., Figs 22a, 46f) and lateral mar- undivided (e.g., Figs 36c, d, 47c, d), usually gins (e.g., Figs 4f, 7f, 8a, 31f, 43a, 46f); para- bifurcate (e.g., Figs 5c, d, 8c, d, 10c, d, 26c, meres moderate size (e.g., Figs 14b, 40b, d, 30c, d, 35, d, 36c, d, 40c, d, 43c, d), 43b) to large (e.g., Figs 36b, 47b), symmet- CAII(L) often basally membraneous, rarely rical, with columnar stem, and weakly (e.g., without lobal sclerites (e.g., Fig. 35c, d), Figs 14b, 26b, 35b, 40b, 43b) to strongly usually with apical lobal sclerite, either hooked (e.g., Figs 8b, 10b, 18b, 22b, 47b) short, subconical (e.g., Figs 10c, d, 26c, d), crown, usually with apex of stem setose, and hooked with spicules (e.g., Fig. 40c, d), or sometimes with flange (e.g., Figs 5b, 8b, digitiform (e.g., Figs 8c, d, 43c, d), some- 35b, 36b); aedeagus with mesial U-shaped times entirely sclerotized (e.g., Figs 22c, d), phallobase, with round capitate processes; sometimes spiculate, CAII(M) usually aedeagus box-like, often heavily sclerotized; membraneous with apical lobal sclerite, ei- phallotheca cylindrical (e.g., Figs 5c, 26c) ther short (e.g., Figs 5c, d), triangular (e.g., to subconical (e.g., Figs 8c, 10c, 22c), thick, Figs 36c, d), digitiform lobal sclerite (e.g., sometimes with pair of subdistal thorn-like Figs 26c, d), hooked (e.g., Figs 40c, d), or processes (Fig. 40c), or areas of sclerotiza- with bifid sclerotized process (e.g., Figs 22c, tion (e.g., Figs 5c, d, 30c, 36c); ductus sem- d, 43c, d); CAIII either separated, heavily inis proximalis narrow, membraneous, with- sclerotized, moderate size, arcuate (e.g. Figs out ribbing (e.g., Figs 5c, 8c, 10c, d, 14c, 10c, d), elongate and sclerotized (e.g., Figs 22c, d, 26c, 30c, 36c, 43c) sometimes thick- 22c, d), medially fused, small (e.g., Figs 8c, er (Fig. 18c), rarely bounded by sclerotized d) or large and U-shaped (e.g., Figs 26c, d, process of vesica (Fig. 30c) or ligamentary 30c, d, 40c, d, 43c, d), or antler-like (Figs tubule (e.g., Figs 18c, 47c, d); ejaculatory 36c, d), or medially fused and membraneous apparatus elaborate (e.g., Figs 5c, 8c, 10c, d, (e.g., Figs 14c, d, 47c, d), sometimes bifur- 14c, 22c, d, 26c, 30c, 36c, 43c), moderately cate, with dorsal and ventral branches (e.g., to heavily sclerotized, most often with con- Figs 5c, d), rarely absent (Fig. 18c). Female voluted ventral conducting canal (e.g., Figs Terminalia: either co-planar or bi-planar 5c, 8c, 10c, d, 22c, d, 26c, 30c, 36c, 43c), (e.g., Figs 33d, f); either ventrally or caudal- with 2-12 convolutions; ejaculatory reser- ly oriented, sometimes recessed; paratergites voir small (e.g., Fig. 22c), moderate size VIII moderate size to large, subtriangular to (e.g., Figs 5c, 8c, 10c, 14c, 43c) to large subelliptoid, usually strongly tapered to- (e.g., Figs 18c, 22c, d, 26c, 30c, 35c, 36c, wards midline; paratergites IX small (e.g., 40c), elliptoid to oval, sometimes greatly re- Fig. 33b) to large (e.g., Figs 33e, f), usually duced and heavily sclerotized (Fig. 47c); ovoid and broader in middle, rarely tapered dorsal conducting canal simple, narrow to medially; gonocoxae I most often entire, relatively broad (e.g., Figs 5c, 8c, 10c, d, rarely subdivided, usually moderate size, 22c, d, 26c, 30c, 36c, 43c); ductus seminis greater than paratergites IX, subtriangular, distalis usually narrow, rarely expanded; posterior margin rectilinear to weakly exca- vesica rarely membraneous (Fig. 18c, d), vate, sometimes depressed medially, and of- usually sclerotized, moderate size (e.g., Figs ten with medial margins raised; first and sec- 5c, d, 22c, d, 36c, d, 47c, d), or elongate ond gonapophyses membraneous; first and (e.g., Figs 8c, d, 10c, d, 26c, d, 30c, d, 35c, second rami usually developed. Spermathe- d, 40c, d, 43c, d), arcuate (e.g., Figs 10c, d, ca: proximal fecundation canal short to

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elongate, membraneous, sometimes strongly denticulate (Figs 11c,d, 12a-c), or body sclerotized basally; reservoir usually broad, very small, < 4 mm (Fig. 24b, 37b) . .(6) oval, often ribbed, rarely weakly dilated, 2 Posterior margin of pronotum strongly membraneous; distal fecundation canal usu- rounded (Fig. 3c-e); pro- and mesosternal ally short, rarely elongate, sometimes heavi- keels overlapping (Fig. 7c); peritreme of ly sclerotized, usually with proximal and dis- metathoracic glands elongate (Fig. 7d) tal flanges, bulb small, oval, sometimes (Coleotichus) ...... 3 bilobed. Larvae: dull or iridescent coloura- – Posterior margin of pronotum rectilinear tion; dorsal abdominal gland openings be- (Fig. 3a), or weakly arcuate, rectilinear tween terga III-IV, IV-V and V-VI. medially (Fig. 3b); pro- and mesosternal Diversity and distribution: The Scutel- keels not overlapping (Figs 4c, 9c); per- leridae are composed of about 80 genera and itreme of metathoracic glands short (Fig. 500 species (LATTIN 1964; Zoological Record 4d) to moderate size (Fig. 9d) ...... 5 1965-2005). Of the scutellerid subfamilies, 3 Moderately sized species, males < 16 mm, the nominotypical Scutellerinae are the females < 17.2 mm; labium reaching at most diverse and are primarily found in the most to metasternum; sub-anterolateral Old World tropics, with only one genus, Au- angles of scutellum without polished or- gocoris BURMEISTER, known from the West- ange callus (Figs 3a, b), at most with a ern Hemisphere. The Pachycorinae are also black spot; scutellum without patterned diverse, with over 100 described species, in- small black spots (Figs 3c, d) ...... 4 cluding numerous undescribed taxa, and are primarily found in the Western Hemisphere, – Large species, males > 20 mm, females > mostly in the Neotropical zoogeographic re- 18 mm; labium reaching abdominal SIV, gion, with a few genera also represented in with medial regions of SII-SIV corre- the Afrotropical (Deroplax MAYR) or spondingly sulcate; sub-anterolateral an- Afrotropical and Oriental regions (Hotea gles of scutellum with polished orange AMYOT &SERVILLE). The Odontotarsinae calli (Fig. 3e); scutellum with pair of are the next most diverse suprageneric black spots sublateral anterior to mid- group, and have putatively the broadest dis- point (Fig. 3e) . . . . . Coleotichus excellens tribution of all the scutellerid subfamilies, 4 Callosite region of pronotum with two with a near cosmopolitan distribution, al- black spots submedially (Fig. 3d); lateral though more than 80 of the species of this margins of posterior 1/2 of scutellum con- subfamily are found in the Palaearctic re- trastingly fuscous (Fig. 3d); males with ab- gion. The Elvisurinae and the Sphaerocori- dominal setose sternal glands on SIV-SVI nae are small subfamilies, with less than 25 (Figs 2a-c); CAII asymmetrical (Figs 8c, species, which are found primarily in the Old d); CAIII medially fused (Fig. 8d); female World tropics. The Eurygastrinae and Tecto- terminalia co-planar; paratergites VIII corinae are currently defined as monogener- large, subelliptoid . . . Coleotichus costatus ic subfamilies, of the Eastern Hemisphere. – Callosite region of pronotum with four black spots, arranged transversely (Fig. Key to Australian Scutelleridae 3c); lateral margins of scutellum concol- orous with remainder of scutellum (Fig. 1 Lateral margins of thoracic sterna strongly 3c); males without abdominal setose ster- sulcate with elevated keels (Figs 4c, 7c, nal glands; CAII symmetrical; CAIII me- 9c); adults dull in colouration (Figs 3a-e), dially separated; female terminalia bi-pla- jugal and pronotal margins entire (Figs nar, paratergites VIII angulate relative to 3a-e) (Elvisurinae) ...... 2 gonocoxae I; paratergites moderately – Lateral margins of thoracic sterna flat, sized, subtriangular . . .Coleotichus artensis without elevated keels (Figs 12c, 20c, 5 Body elongate-oval (Fig. 3a); dark brown 29c, 31c, 39c, 43c, 46c); adults brightly species with heavy punctation (Fig. 3a); coloured (Figs 1a-f, 16a, b, 19a-d, 24a,c-f, ventral surface strongly setose; labium 28a-e, 45a, b), or if dull in colouration reaching posterior margin of abdominal SI- (Figs 14a-c), jugal and pronotal margins II abdominal; males without abdominal

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sternal glands; connexival margins thick- (Figs 11a-c); dorsum densely setose, with ened, each segment bicoloured yellow then curly sericeous setae (Figs 13a, b, e, h) in- black (Fig. 3a); CAIII bifurcate, medially termixed with fan-like (Figs 13c, h, i) or fused (Fig. 4d); spermathecal reservoir clove-like setae (Figs 13b, f); male ab- weakly dilated . . . . . Austrotichus rugosus dominal SVIII recessed, not visible (Fig. – Body ovoid (Fig. 3b); orange-brown 12e); female gonocoxae I tripartite species with shallow punctation (Fig. 3b); (Odontotarsinae: Morbora) ...... 9 ventral surface mostly glabrous; labium – Body large, males and female > 13 mm; reaching metasternum; males with ab- body mostly orange, most often with iri- dominal sternal glands on SIV-SVII con- descent blue or green markings, rarely nexival margins greatly thickened, con- uniformly orange (Figs 1f, 45a, b; dorsum colorous; CAIII not bifid, medially sepa- glabrous; male abdominal SVIII exposed, rated (Figs 10c, d); spermathecal reservoir broadly covering ventral surface of py- broad, oval . . . . . Solenotichus circuliferus gophore (Figs 45c, 46e); female gonocox- 6 Both sexes with stridulatory vittae on ab- ae I undivided . . . Tectocoris diophthalmus dominal SV-SVI (Fig. 17e); head large 9 Anterolateral margins of pronotum and (Figs 16a, b); SVII enlarged, covering male costal margins greatly expanded (Figs (Fig. 17d) and female terminalia (Pachy- 11c); scutellum with broad fan-like setae corinae) ...... Agonosoma trilineatum (Figs 13h-i) ...... Morbora schoutedeni – Both sexes without stridulatory vittae on – Anterolateral margins of pronotum and abdominal SV-SVI; head moderately sized costal margins not expanded (Fig. 13a, b); (e.g., Figs. 19a-d); SVII not greatly en- scutellum with either clove-like setae larged (e.g., Figs 21a-i), terminalia of both (Figs 13b, 13f) or narrow fan-like setae sexes exposed, sometimes male SVIII ex- (Fig. 13c) ...... 10 posed partially covering pygophore (e.g., 10 Dorsum with short, narrow fan-like setae, Figs 45c, 46f) ...... 7 strongly tapered proximally (Fig. 13c); AI 7 Males with androconial glands on abdom- longest antennal segment; CAI branched; inal SIV-VI (Figs 2d-f, g-i, 12e, 45c); ex- CAIII absent ...... Morbora australis ternal efferent system of metathoracic – Dorsum without fan-like setae, with glands reduced (e.g., Figs 12d, 46d); clove-like setae only (Fig. 13f); AIV aedeagal ejaculatory apparatus without longest antennal segment; CAI branched; convoluted ventral conducting canal CAIII present ...... Morbora hirtula (Figs 14c, d, 47c, d); CAI present (Figs 14c, d, 47c, d); CAIII well-developed, 11 Body very small, < 4 mm (Figs 24b, 37); membraneous, basally fused (Figs 14c, d, body with setigerous punctures (Figs 38a- 47c, d) ...... 8 c); head short, strongly rounded in profile (Fig. 38b); jugal margins rounded (Fig. – Males without androconial glands, most 38b); apex of peritreme of metathoracic often with abdominal sterna unadorned glands distally raised, obovate (Fig. 38c) (e.g., Figs 21a-i); external efferent system ...... Heissiphara minuta of metathoracic glands usually well-devel- oped (e.g., Figs 20d, 31d), if evaporative – Body small to large species, > 8 mm; body areas reduced, peritreme greatly enlarged, without setigerous punctures, dorsum usu- subtriangular (Figs 25d, 29d); aedeagal ally glabrous, at most with simple setae; ejaculatory apparatus with convoluted head short to elongate; jugal margins car- ventral conducting canal (e.g., Figs 5c, 8c, inate (e.g., Figs 29b, 31a, b) or rounded 10c, d, 22c, d, 26c, 30c, 36c, 43c); CAIII (e.g., Figs 25c); peritreme subreniform usually large, membraneous and U-shaped (e.g., Figs 20d, 39d, 42d), subtriangular (e.g., Figs 26c, d, 30c, d, 40c, d, 43c, d), (e.g., Figs 25d, 29d), rarely obovate, but if rarely antler-like (Figs 36c, d) or digiti- so anteriorly directed (Fig. 31d) . . . . . 12 form (Fig. 22c, d) (Scutellerinae) . . . . 11 12 Jugal margins straight in dorsal view 8 Body small, males and females < 6 mm; (Figs 28a, 29a); large orange species with body dull colouration, densely punctate seven black spots on pronotum and seven

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spots on scutellum (Fig. 28a); abdominal – Dorsum mostly iridescent green (Fig. 28e) SVIII exposed (Figs 21e, 29e), covering or blue, often with anterior orange callus anterior margin of pygophore; CAII undi- on scutellum (Fig. 1d); abdominal venter vided (Figs 30c, d); female terminalia cau- mostly iridescent dark green or blue, with dal in orientation . . . . . Cantao parentum orange lateral margins, inner outline of – Jugal margins sinuate in dorsal view (e.g., latter notched (Fig. 21h); anterior callus Figs 20a, 25a, 31a, 39a, 42a); various of scutellum impunctate (Figs 1f, 28e); colouring, if orange never with above- CAII(L) apex linear ...... mentioned black markings; male abdomi- ...... Lampromicra senator nal SVIII recessed, not visible; CAII di- 16 AII(b) minute, shortest antennal seg- vided (e.g., Figs 22c, d, 26c, d, 35c, d, 40c, ment; AII(a) arcuate, more so in males; d, 43c, d); female terminalia caudal or dorsum strongly iridescent green with or- ventral in orientation ...... 13 ange and fuscous markings (Fig. 28b); lat- 13 Dorsum densely setose (Fig. 1d); body iri- eral areas of mesosternum striate; crown descent copper-dark brown (Fig. 28c), of paramere weakly curved, apex truncate green (Fig. 1f), bicoloured dark- (Fig. 43b); CAII(L) with an elongate ar- green/purple and yellow (Fig. 28d) or blue cuate lobal sclerite (Figs 43c, d) ...... (Fig. 28e), often with contrasting coloura- ...... Scutiphora pedicellata tion (e.g., Figs 1d, 28d); CAII(L) with – AI or AII(a) shortest segment; AII(a) lin- perpendicular or hook-shaped apex, dis- ear; body usually red or orange with con- tally denticulate (Fig. 40c, d) (Lampromi- trasting markings, if iridescent green cra) ...... 14 without orange markings; lateral areas of – Dorsum usually glabrous, at most with mesosternum smooth; crown of paramere scattered setae; CAII(L) either with digi- strongly hooked (e.g., Fig. 36b); CAII ei- tiform (Fig. 22c, d) or subconical lobal ther undivided, or with base membrane- sclerite (Fig. 26c, d), or bag-like (Figs 35c, ous post-thecal margin (e.g., Figs 35c, d, d, 36c, d), with or without basal denticu- 36c, d), if entirely sclerotized, oval in lation ...... 16 shape ...... 17 14 Posterior 1/2 of scutellum yellow, re- 17 Body small to moderate size, males < 12.5 mainder iridescent dark-green/purple mm, females < 13 mm; dorsum densely (Fig. 28d); abdominal venter mostly iri- punctate (Figs 24c-f); jugal margins cari- descent green, with yellow lateral mar- nate (Figs 31a, b); hemelytral membrane gins, inner outline of latter linear (Fig. tip not exposed beyond scutellum (Figs 21g); CAIII medially fused post-thecal 24c-f); peritreme of metathoracic glands margin ...... obovate, elongate, laterally recurved to- ...... Lampromicra regia wards head (Fig. 31d); ventral surface of male pygophore caudally oriented (Figs – Posterior 1/2 of scutellum either copper- 31e, 32b, f, h) (Choerocoris) ...... 18 fuscous (Fig. 28c) or iridescent dark green or blue (Fig. 1f, 28e); abdominal venter – Body moderate size to large, males > 13 with margins of lateral colour band linear mm, females > 13.5 mm; dorsum punctate (Fig. 21f) or notched (Fig. 21h); CAIII to impunctate; hemelytral membrane tip separated post-thecal margin (Fig. 40d) exposed beyond scutellum (Figs 19a-d); 15 peritreme of metathoracic glands either 15 Dorsum uniformly copper-fuscous, some- subtriangular (Fig. 25e) or subreniform times with iridescent green or purple (Fig. 20d); ventral surface of male py- sheen (Fig. 28c); abdominal venter most- gophore ventrally oriented (Figs 20 a-c) ly concolorous with dorsum, with red lat- or caudally oriented ...... 21 eral margins, inner outline of latter linear 18 Callosite region of pronotum medially (Fig. 21f); anterior callus of scutellum impunctate (Fig. 24d); callosite region de- punctate, always fuscous (Fig. 28c); marcated posteriorly by transverse furrow CAII(L) apex hook-like (Figs 40c, d) . . . (Fig. 24d); scutellum red with pair of sub- ...... Lampromicra aerea medial, iridescent blue, subtriangular

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markings (Fig. 24d), anterior margin with gonocoxae I (Figs 33c, d); gonocoxae I iridescent blue marking; apex of scutel- well-developed, posterior margin weakly lum without markings (Fig. 24d); abdom- excavate (Figs 33c, d) ...... inal venter mostly red with lateral and ...... Choerocoris variegatus submedial dark blue markings (Figs 32e, 21 Hemelytral membrane barely surpassing f); CAII(M) and CAII(L) roughly sym- tip of scutellum (Fig. 24a); dorsum mostly metrical ...... Choerocoris paganus fuscous, with orange/red and iridescent – Pronotum densely and uniformly punc- dark green markings (Figs 24a); AII(a) tate (Figs 24c, e, f); callosite and disc re- moderately short, AII(b) at most 2.5x > gions of pronotum co-planar (Figs 24c, e, AII(a); peritreme of metathoracic glands f); scutellum orange or red with three greatly expanded laterally, subtriangular, prominent iridescent blue regions, always 1-1.5x longer than wide (Figs 25d); evap- with V-shaped distal markings (Figs 24c, orative areas restricted, not extending to e, f); abdominal venter mostly orange or mesepimeron (Fig. 25d); CAI absent (Figs red, with (Figs 32a, b) or without subme- 26c); female terminalia caudal in orienta- dial darker markings (Figs 32c, d, g, h); tion ...... Calliscyta stalii CAII(M) and CAII(L) significantly dif- – Hemelytral membrane clearly surpassing ferentiated (Figs 35c, d, 36c, d) . . . . . 19 tip of scutellum (Figs. 19a-d); dorsum 19 Jugae orange (Fig 24c); abdominal venter mostly iridescent green, orange, or orange mostly orange with lateral blue or green and blue, with various markings (Figs markings (Figs 32c, d); apex of CAII(M) 19a-d); AII(a) minute, AII(b) more than with weak sclerotization (Fig. 35c, d); 6x > AII(a) length; peritreme of metatho- CAIII contiguous medially, lightly sclero- racic glands subreniform, tapered distally, tized, lobe-like (Figs 35c, d); vesica with- 3x or more longer than wide (Fig. 20d); out distal process beyond secondary gono- evaporative areas extensive, extending to pore (Fig. 35c, d) ...... mesepimeron (Fig. 20d); CAI present ...... Choerocoris lattini nov.sp. (Figs 22c, d); female terminalia ventral in – Head mostly iridescent dark green or blue orientation (Calliphara) ...... 22 (Figs 24e, f); abdominal venter either yel- 22 Eye length 1/2 head length (Fig. 19d); low or orange, with (Fig. 32a, b) or with- head orange-red to copper with fuscous out submedial dark (Figs 32g, h) blue markings (Fig. 19d); pronotum iridescent markings; apex of CAIII with prominent copper, with calli most often fuscous (Fig. triangular lobal sclerite (Figs 36c, d); 19d); scutellum iridescent copper, with CAIII separated, as antler-like and heavi- fuscous spot at tip (Fig. 19d); exocorium ly sclerotized processes (Figs 36c, d); vesi- iridescent green (Fig. 19d); posterior an- ca with subdistal paired process (Figs 36c, gles of abdominal SVI-SVII spinose; CAI d) ...... 20 bifurcate, with CAI(D) elongate, digiti- 20 Pregenital abdominal venter mostly yel- form (Fig. 22c, d); CAIII divided (Figs low with submedial and lateral black 22c, d); vesica with apical hood (Fig. 22c, bands (Figs 32a,b); posterior angles of ab- d) ...... Calliphara regalis dominal connexiva V-VII nodulate, yel- – Eye length < 1/2 head length (Figs 19a- low (Figs 32a, b); female paratergites IX c); head concolorous, iridescent dark greatly expanded, much larger than gono- green or blue; pronotum orange or irides- coxae I (Figs 33e, f); gonocoxae I greatly cent green, rarely with calli fuscous; reduced, posterior margin deeply excavate scutellum broadly bicoloured (Fig. 19a), (Figs 33e, f) . . . Choerocoris grossi nov.sp. or orange with narrow distal iridescent – Pregenital abdominal venter mostly yel- green marking (Fig. 19b); posterior an- low with lateral black band only (Figs gles of abdominal sterna without spines 32g, h); posterior angles of abdominal (Fig. 21c), or SIV-SVI (Fig. 21b) or SIV- connexiva V-VII weakly swollen at most, SVII spinose (Fig. 21a); CAI and CAIII fuscous (Figs 32g, h); female paratergites divided or undivided; vesica with or IX not greatly enlarged, smaller than without processes ...... 23

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23 Scutellum either red and iridescent dark green iridescence; larvae brilliantly and blue (Fig. 19a) or mostly orange (Fig. aposematically coloured; head subtriangular 19b); legs concolorous, fuscous; abdominal to triangular (e.g., Figs 4a, 7a, 9a); sulcate venter impunctate (Figs 21a, b), some- thoracic sterna with highly elevated keels times rugose laterally; spinose posterolat- (e.g., Figs 4c, 7c, 9c), contiguous with ex- eral angles of abdominal sterna SIV-SVI planate anterior margin of proepisternum; (Fig. 21b) or SIV-SVII (Fig. 21a) . . . . 24 efferent system of metathoracic glands well- – Scutellum mostly iridescent green, each developed (e.g., Figs 4d, 7d, 9d), with per- with seven black spots (Fig. 19c); femora itreme short (e.g., Fig. 4d) to elongate (e.g., mostly orange, apex narrowly fuscous; tib- Fig. 7d); sometimes males with abdominal iae and tarsi fuscous; abdominal venter sternal glands (e.g., Figs 2a-c); ventral sur- punctate (Fig. 21c); posterior angles of face of pygophore caudally (e.g., Fig. 4e) or abdominal sterna not spinose (Fig. 19c) . ventrally oriented (e.g., Fig. 7e); posterior ...... Calliphara nobilis margin of male pygophore emarginate (e.g., Figs 4e, 7e) to entire (e.g., Fig. 9e); genital 24 Scutellum mostly orange, with extreme opening narrow to broad; parameres hook- caudal tip iridescent green (Figs 1a, 19b); shaped (e.g., Figs 5b, 8b, 10b); ejaculatory dorsum with moderately deep punctation; reservoir well-developed, with ventral con- abdominal pregenital venter mostly iri- ducting canal and ejaculatory apparatus descent green, laterally red to orange-red, (e.g., Figs 5c, d, 8c, d, 10c, d); CAI absent; rugose (Fig. 21b); posterolateral angles of CAII bifurcate, with lobal sclerites (e.g., abdominal SIV-SVI spinose (Fig. 21b) . . Figs 5c, d, 8c, d, 10c, d), CAIII undivided ...... Calliphara imperialis (e.g., Figs 8c, d) or divided (e.g., Figs 5c, d, – Scutellum broadly bicoloured, anterior 10c, d); spermathecal fecundation canal 1/2 orange to orange-red, remainder iri- short or long; and, spermathecal reservoir descent dark blue, with w-shaped anterior oval to weakly dilated. outline (Fig. 19a); dorsum with shallow Remarks: STÅL (1872) first described the punctation; abdominal pregenital venter elvisurines as a tribal group (Elvisuraria), pro- orange, laterally smooth (Fig. 21a); pos- viding a diagnosis (STÅL 1872, 1873) based terolateral angles of abdominal SIV-SVII primarily on the carinate thoracic sterna. spinose (Fig. 21a) ...... SCHOUTEDEN (1904) followed this arrange- ...... Calliphara dimidiata cruenta ment and provided a listing of species. Subfamily Elvisurinae KIRKALDY (1909) catalogued the species and

Pentatomidae, Scutellerina, Elvisuraria STÅL the classification of SCHOUTEDEN (1904). 1872: 32 (new suprageneric taxon) The Elvisurinae are retained as a subfam- Pentatomidae, Scutellerinae, Elvisuraria: ily in this work, on the basis of the keel-like SCHOUTEDEN 1904: 4 (diagnosis) thoracic sterna (Figs 4c, 7c, 10c), which is a Pentatomidae, Scutellerinae, Elvisurini: putative apomorphy for this taxon. In some KIRKALDY 1909: 311 (catalogue) Pentatomidae, Scutellerinae, Scutellerini, Elvi- Odontotarsinae species, the pro- and suraria: LESTON 1952d: 13 (new suprageneric mesosterna are raised, but not to any great classification) height, and not keel-like as in the elvisurines. Scutelleridae, Elvisurinae: MCDONALD & CASSIS Unlike, the Scutellerinae and some 1984: 538 (Australia) Pachycorinae, the adults of elvisurines are Diagnosis: The Elvisurinae are recog- nearly all dull in colouration, aside from two nised by the following combination charac- species of Coleotichus (C. blackburniae and C. ters: small (e.g., Fig. 3b) to very large species bulowi), which have a green iridescence, and (e.g., Fig. 3e); ovoid to elongate-ovoid, some species of Chiastosternum and Solenos- sometimes strongly tapered towards termi- thedium, which have contrasting yellow nalia; body moderately to heavily punctate; markings on the dorsum. It is noteworthy pale to dark brown species (e.g., Figs 3a-e), that larval elvisurines have brilliant coloura- sometimes with red or fuscous highlighting tion, like the larvae in the Scutellerinae. In (e.g., Fig. 3d), occasionally with patterning most other characters, the elvisurines are in- on scutellum (e.g., Fig. 3d), sometimes with distinguishable from the other jewel bug sub-

300 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at families, aside from the Tectocorinae. The male aedeagus of elvisurines is not diagnostic for the subfamily, and closely resembles the aedeagus in taxa of the other subfamilies, particularly the Scutellerinae (e.g., Choero- coris species, Figs 35c, d, 36c, d), Pachycori- nae (e.g., DALLAS; MCDONALD 1966: Fig. 30) and Odontotarsinae (e.g., Phi- modera GERMAR; MCDONALD 1966: Fig. 22); with a well-developed ejaculatory apparatus, including a convoluted ventral conducting canal and a prominent ejaculatory reservoir. In addition, the character states of the con- junctival appendages are difficult to inter- pret, with the CAI absent in elvisurines, but also in many taxa in all subfamilies except the Tectocorinae. Moreover, the female ter- minalia and spermatheca of elvisurines is consistent with most other scutellerids. At present, there is no conclusive evidence that suggests a sister-taxon relationship with any of the other scutellerid subfamilies. The Elvisurinae are a small subfamily of scutellerids comprised of five genera (KIRKALDY 1909; LESTON 1952d). This sub- family is restricted to the Eastern Hemi- sphere, with the genera distributed as fol- lows: the monotypic genera Austrotichus and Solenotichus are endemic to Australia; Solenosthedium has seven species in the East- ern Hemisphere (from to Indonesia, including Madagascar), aside from the Aus- tralian zoogeographic region; Coleotichus is broadly distributed in the Australian region, including continental Australia and the Melanesian subregion, with spectacular species in Samoa (C. bulowi) and (C. blackburniae WHITE); the nominotypical genus Elvisura is endemic to the southern re- gions of the Afrotropical region, including Madagascar, to (KIRKALDY 1909). The genus Nesogenes, a monotypic genus (N. boscii FABRICIUS), was described by HORVÁTH (1921), and is known from the Greater Antilles (Cuba, Haiti and St. Vin- cent). He placed this taxon in the Elvisuri- nae, based on the conformity of the keel- Fig. 3: Habitus of Australian like thoracic sterna with all the other mem- Elvisurinae species bers of the subfamily. We examined a single (a) Austrotichus rugosus, = specimen of this taxon at the BMNH (label (b) Solenotichus circuliferus, = data as ‘St. Dom.’), and found that the (c) Coleotichus artensis, = (d) Coleotichus costatus, = species has elvisurine-type thoracic sterna, (e) Coleotichus excellens, =. but possesses extensive stridulatory vittae Scale bars: 1 mm.

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(sublaterally on abdominal SIII-SVII), the panded anteriorly (Fig. 4c). Pronotum: latter being diagnostic for the Pachycorinae. strongly transverse, subtrapezoidal; anterior On these two character systems alone, this margin weakly concave (Fig. 3a); anterolat- taxon is annectant between the Elvisurinae eral margins subcarinate; posterior margin and the Pachycorinae. Externally, this rectilinear (Fig. 3a); callosite region weakly species is reminiscent of species of demarcated; disc strongly convex, declivent Coleotichus, having a rounded posterior mar- posteriorly (Fig. 3a). Scutellum: broad, gin of the pronotum, and densely punctate strongly depressed beyond anterior 1/2; el- dorsum, with green iridescence (as in C. liptoid tumescence on anterior margin, blackburniae and C. bulowi). The external marked sublaterally by dark foveae (Fig. 3a); efferent system of metathoracic glands is ex- lateral margins convex (Fig. 3a); base of tensive, with no peritreme evident, but with corium and clavus, and connexiva III-VII a prominent ostiole; a character state con- exposed (Fig. 3a). Thoracic pleura: external forming to many Western Hemisphere efferent system of metathoracic glands well- pachycorine taxa. The external female gen- developed (Fig. 4d); evaporative areas occu- italia are distinctive, with the medioposteri- pying most of metepisternum, extending to or angles of the first gonocoxae narrowly ex- mesepimeron (Fig. 4d); ostiole small (Fig. cavate. There has been no examination of 4d); peritreme short, raised, reniform, not the male genitalia of this species. We regard medially sulcate (Fig. 4d). Thoracic sterna: this taxon as incertae sedis, pending exami- lateral margins sulcate, prosternal and nation of additional material, but exclude it mesosternal keels not overlapping (Fig. 4c); from the Elvisurinae. anterior margin of proepisternum truncate, not elevated (Fig. 4c). Male Genitalia: py- Austrotichus GROSS 1975 gophore (Figs 4e, f, 5a) caudally oriented, (Figs 3a, 4, 5, 6) moderately large, transverse; genital open- Austrotichus GROSS 1975: 83 (gen. nov.); MC- ing dorsal, narrow, suboval, lateral margins DONALD & CASSIS 1984: 538, 545 (key descrip- setose (Figs 4f, 5a); parameres with colum- tion, genitalia); CASSIS & GROSS 2002: 582 (cat- nar stem, moderately hook-shaped crown, alogue) flange at base of crown (Fig. 5b); aedeagus Type species: Austrotichus rugosus GROSS 1975, by original designation (Figs 5c, d): phallotheca short, without processes; ductus seminis proximalis narrow, Diagnosis: Austrotichus is recognised by singular, membraneous, tubelike; ejaculato- the following characters: posterior margin of ry apparatus moderately developed; ventral pronotum rectilinear (Fig. 3a); thoracic conducting canal with about eight paired sterna carinate, not overlapping (Fig. 4c); convolutions, heavily sclerotized; ejaculato- anterior margin of proepisternum truncate ry reservoir small, subelliptical; dorsal con- (Fig. 4c); peritreme of metathoracic glands ducting canal narrow; ductus seminis distal- short, arcuate (Fig. 4d); posterior margin of is and vesica basally incrassate, sinuate, api- male pygophore medially emarginate (Fig. cally truncate; CAI absent; paired CAII 4e); CAI absent (Figs 5c, d); CAII(L) with- greatly enlarged, membraneous, distally bi- out lobal sclerite (Figs 5c, d); CAII(M) with furcate, lateral branch without sclerotiza- short lobal sclerite (Figs 5c, d); CAIII bifid, tion, medial branch with small, acute lobal medially fused, CAIII(V) with short lobal sclerite (Figs 5c, d); paired CAIII mostly sclerite (Figs 5c, d); gonocoxae I not divid- membraneous, with lateral margins sclero- ed, medial margins raised; and, spermathe- tized, bifurcate, lateral branch membrane- cal reservoir membraneous, weakly dilated, ous, medial branch with small acute lobal elliptoid. sclerite, medial pair fused along midline Description: Body elongate-ovoid (Fig. (Figs 5c, d). Female Abdominal Venter: pos- 3a); dorsum and venter strongly convex; terior margin of SVII weakly bisinuate, wing tip not extending beyond scutellum weakly thickened medially. Female Termi- (Fig. 3a). Head: mostly flattened, posterior nalia: caudally oriented, bi-planar; parater- aspect of vertex weakly convex (Fig. 4a); ju- gites VIII subelliptoid, medially raised; gal margins thickened, subcarinate (Figs 4a, paratergites IX small, narrowly elliptoid, b); bucculae margins subparallel, weakly ex- medially rounded; gonocoxae I moderately

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Fig. 4: Scanning electron micrographs of key characters of Austrotichus rugosus (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: C = clypeus; Ea = evaporative area; Go = genital opening; J = juga; Me = mesepi- meron; Met = metepister- num; Ms(k) = mesosternal keel; Pe = peritreme; Pe(k) = proepisternal keel, ventral view; P(r) = right paramere; Pr = proctiger. Scale bars = 1 mm.

large, medially depressed, posterior margin Remarks: GROSS (1975) described Aus- concave, medial margins elevated. Sper- trotichus as a monotypic genus, based on ex- matheca: short proximal fecundation canal; ternal characters alone. MCDONALD & CAS- spermathecal reservoir narrowly dilated, el- SIS (1984) supported its generic status, and liptoid, membraneous, without pronounced described the male and female genitalia. It is modifications; pump short, with dorsal and a distinctive genus of jewel bug, with a ro- bust body, apomorphic male genitalia and proximal flanges; bulb oval. metathoracic glands, and specialized host Diversity and distribution: Austrotichus is plant associations. Unlike species of a monotypic genus, which is broadly distrib- Coleotichus, this genus has the posterior mar- uted in temperate Australia, including Victo- gin of the pronotum rectilinear (Fig. 3a) (al- ria, and Western Australia. so in Solenotichus and Afrotropical Solenos- thedium SPINOLA) and the anterior margin of Included species: the scutellum is moderately tumose, with A. rugosus GROSS 1975 Australia sublateral dark foveae (Fig. 3a). The male

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Fig. 5: Male genitalia of Austrotichus rugosus (a) pygophore, dorsal view (b) right paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII(D) = third conjunctival appendage, dorsal branch; CAIII(V) = third conjunctival appendage, ventral branch; DCC = dorsal conducting canal; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; F = flange at base of crown of paramere; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

304 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at genitalia of Austrotichus rugosus, with the paired CAIII mostly membraneous (Fig. 5c, d), differs from species of Coleotichus, Solenotichus circuliferus, and extralimital species of Elvisurinae, where the CAIII is sclerotized. In addition, the peritreme is short in Austrotichus (Fig. 4d), much shorter than in Solenotichus (Fig. 9d), still apparent in comparison to Solenosthedium, and much shorter than the consistently elongate per- itreme of species of Coleotichus (Fig. 7d).

Austrotichus rugosus GROSS 1975 (Figs 3a, 4, 5, 6, Table 4)

Austrotichus rugosus GROSS 1975: 582 (n.sp.); MCDONALD & CASSIS 1984: 544, Figs 17-22 (de- scription, = & Y genitalic illustrations); CASSIS & GROSS 2002: 582 (catalogue) Diagnosis: Austrotichus rugosus is recog- nised by the following combination of char- acters: body strongly convex dorsally; dark grey-brown (Fig. 3a); heavily punctate (Fig. 3a); AII(a) short; AIV longest segment; Structure. Antennae: AI relatively Fig. 6: Distribution of Australian labium extending to posterior margin of ab- elongate, just surpassing lateral margins of Elvisurinae species. dominal sterna IV; and, genitalia as in head; AII(a) short; AII(b) and AIV roughly generic description (Fig. 5). subequal in length; AIII longest segment. Labium: reaching between posterior margin Description: Body moderately large, males of abdominal sternite IV; LII longest seg- 10.03-12.22 mm, females 11.10-12.66 mm. ment; LIV shortest segment. Abdominal Colouration. Body mottled yellow- Venter: posterior angles of SIV-VII expand- brown to most often dark grey-brown (Fig. ed (Fig. 3a). Male genitalia as in generic de- 3a), punctures black, with scattered polished scription (Fig. 5). yellow regions on callosite region of prono- tum. Head: darker brown on clypeal margins Measurements. Table 2. MCDONALD & extending to posterior margin of vertex; an- CASSIS (1984) did not provide measure- tennae dark-brown, sometimes AI proximal- ments for this species. ly yellow; labium uniformly fuscous. Prono- Type material examined: Holotype: =, Port Lin- tum: calli with dark brown outline. Scutel- coln, South Australia (SAMA I20,400). lum: anterolateral foveae black. Thoracic Paratypes: 1Y, same data as holotype (SAMA pleura: yellow-brown. Pregenital Abdomen: I20,402); 1Y, Keith, South Australia, ex leaf lit- venter mostly mottled yellow-brown, spirac- ter, June 1952, MG coll. (SAMA I20, 404; Y ular-trichobothrial region polished yellow; paratype seen); 1 , 23 mi. E Ravensthorpe, = posterior angles of SIV-VIII with anterior Western Australia (SAMA I20,600); 1 , Wemb- ley Park, 8 mi. N Perth, Western Australia, 4-xi- 1/2 yellow, posterior 1/2 fuscous (Fig. 3a). 1935, RE Turner, BM 1935-240 (BMNH); 1Y, Texture. Body densely punctate, punc- Dedari, 40 mi. W Coolgardie, ii-21-i-1936, RE tures irregularly distributed often coalesced. Turner BM 1936-28 (BMNH). Other material examined: : 1Y, Little Vestiture. Dorsum almost glabrous; an- Desert National Park, Stans Camp Track, tennae: AI almost glabrous, sometimes 36°35.211’S 138°33.732’E, 173 m, RT Schuh, G AII(a,b) also glabrous or nearly so, AIII- Cassis, MD Schwartz & R Silveira, 6 November AIV with moderate distribution of short 2002, ex Allcasuarina pusilla (AM); Southern semierect setae; genae pilose; abdominal Australia: 1Y, Springs Conservation venter moderately pilose, more so on lateral Park, 35.626S 140.867E 100m, G Cassis, RT margins, sternal sutures and terminalia. Schuh and R Silveira, 10 November 1998, site

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Table 4: Austrotichus rugosus: diagnostic measurements in millimetres. N = sample size. feeding behaviour, although their presence Male Female on seed cones is suggestive of seed-preda- N Mean±SD Range N Mean±SD Range tion, as opposed to -feeding proposed by Length 9 11.08±0.59 10.03-12.22 4 12.03±0.66 11.10-12.66 CASSIS & GROSS (2002). GROSS (1975) in Pronotal width 9 7.65±0.36 6.96-8.23 4 8.32±0.70 7.35-8.96 describing this species, documented that Width between eyes 9 2.39±0.19 2.14-2.78 4 2.51±0.15 2.34-2.63 one of the paratypes was found in leaf litter. Antennal segment length This is consistent with other jewel bugs, I 17 1.04±0.08 0.93-1.23 8 1.19±0.06 1.10-1.27 such as Choerocoris paganus, which are II(a) 17 0.59±0.05 0.51-0.69 8 0.66±0.04 0.59-0.71 II(b) 16 1.23±0.07 1.10-1.35 8 1.25±0.09 1.13-1.40 known to be pre- and post-dispersal seed III 7 1.38±0.07 1.27-1.47 7 1.32±0.06 1.25-1.40 predators. IV 3 1.25 1.18-1.35 4 1.26±0.01 1.25-1.27 Labial segment length Remarks: GROSS (1975) described this I 9 1.92±0.06 1.84-2.01 4 1.90±0.02 1.86-1.91 species from a handful of specimens from II 9 2.08±0.13 1.96-2.25 4 2.06±0.07 1.96-2.13 South Australia and Western Australia. He III 9 1.38±0.11 1.23-1.52 4 1.43±0.06 1.35-1.47 IV 9 1.26±0.11 1.18-1.54 4 1.27±0.05 1.23-1.35 also provided a habitus illustration and de- tailed description of non-genitalic characters. 98-40, ex Allocasuarina pusilla (AM); Western MCDONALD & CASSIS (1984) described the == YY Australia: 3 5 , 17 km W of Brand High- male and female genitalia and first recognised way on Green Head Road, 30.05S, 115.133E, that the peritreme of the metathoracic glands 350m, G Cassis and RT Schuh, 1 November is distinctively short. The male genitalia are 1996, site 96-53, ex Allocasuarina humilis (AM); largely invariant across its distributional 1Y, Madfish Bay, Bay National Park, 35.017S, 117.25E, 100 m, G Cassis, RT Schuh range. and R Silveira, 1 December 1999, site 99-54, ex Allocasuarina humilis (AM); 2YY, Eneabba, RP Coleotichus WHITE 1839 McMillan, 12-viii-1978 (WAM). (Figs 3c-e, 6, 7, 8) Distribution: Austrotichus rugosus is Coleotichus WHITE 1839: 541 (nov. gen.); DALLAS broadly distributed in temperate Australia, 1851: 5 (description); STÅL 1865: 35 (key); MAYR from Western Australia to Victoria, includ- 1866: 13 (diagnosis); STÅL 1873: 4 (diagnosis); LETHIERRY & SEVERIN 1893: 15 (catalogue); ing South Australia (Fig. 6). This work in- SCHOUTEDEN 1904: 4, 5 (description); KIRKALDY cludes new records from Victoria, South 1902: 172 (Hawaii); SCHOUTEDEN 1905 (mono- Australia and Western Australia. The graph); KIRKALDY 1909: 313 (catalogue); GROSS species exhibits a significant distributional 1975: 80 (description); DISTANT 1920: 143 (cata- disjunction, between southwest Western logue); GROSS 1975: 80 (description); MCDON- Australia and the southern region of the ALD &CASSIS 1984: 538 (genitalia); CASSIS & Murray-Darling basin. GROSS 2002: 583 (catalogue) Type species: costatus FABRICIUS 1787 by Host plants and biology: The Green monotypy Head Road specimens of Austrotichus rugo- sus were found on the sheoak species, Allo- Diagnosis: Coleotichus is recognised by humilis (Casuarinaceae); a monoe- the following combination of characters: cious commonly found in heath habi- elongate-ovoid (Figs 3c-e); lateral margins of tats in southwest Western Australia. Both jugae rounded (Fig. 7b); posterior margin of sexes were collected on the female seed pronotum strongly convex (Figs 3c-e), cov- cones, in association with the true bug ering anterior margin of scutellum; metatho- racic peritreme mostly rectilinear, sulcate, species Laryngodus australis HERRICH-SCHA- with apex anteriorly arcuate (Fig. 7d); tho- EFFER (: ) and racic sterna strongly keel-like, pro- and (WESTWOOD) (Pentato- midae). Despite their large size, these jewel mesosternal keels overlapping (Fig. 7c); CAI bugs are surprisingly cryptozoic, with their absent; CAII symmetrical or asymmetrical, bifid (Figs 8c, d), with rounded or digitiform mottled colouration blending closely with lobal sclerites; CAIII separated or fused (Fig. the dark brown colour of the female cones. 8d); and, spermathecal reservoir oval. Similarly, the South Australian specimen was collected on the female seed cones of Description: Body elongate-ovoid (Figs another species of Allocasuarina, A. pusilla. 3c-e); moderately-sized to large species; We have not made direct observations of moderately convex dorsally and ventrally;

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Fig. 7: Scanning electron micrographs of key characters of Coleotichus costatus (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ea = evaporative area; Ms(k) = mesosternal keel; Pe = peritreme; Pe(k) = proepisternal keel, ventral view; EES = external efferent system. Scale bars = 1 mm.

wing tip extending beyond scutellum; mod- metasternum (Fig. 7c) to abdominal SIV; erately to densely punctate; pale to dark LII usually longest segment. Pronotum: brown, sometimes with red or green irides- large, shield-like, strongly convex (Figs 3c- cent highlighting (e.g. Coleotichus blackbur- e); anterior margin concave; anterolateral niae; Hawaii), rarely with patterned darker margins rectilinear, carinate; humeral angles brown markings. Head: triangular (Figs 3c- rounded to weakly angulate; posterior re- e, 7a), moderately convex; clypeus broadly gion of disc greatly expanded, posterior mar- rounded at apex and barely surpassing juga gin strongly convex, overlapping anterior (Fig. 7a); jugal margins rounded (Fig 7b); lo- margin of scutellum (Figs 3c-e). Scutellum: rae strongly demarcated, margins carinate elongate, strongly convex, U-shaped, most (Fig. 7b); bucculae narrow, lateral margins often strongly tapered posteriorly (Figs 3c- weakly arcuate (Fig. 7c). Antennae: AI- e); basal aspects of corium and clavus, and AII(a) shortest segments, subequal; AIII- connexiva III-VII exposed (Fig. 3c-e). Tho- AIV longest segments, subequal or AIV racic pleura: external efferent system of slightly longer. Labium: reaching apex of metathoracic glands well-developed, raised

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Fig. 8: Male genitalia of Coleotichus costatus (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L)L = left second conjunctival appendage, lateral branch; CAII(L)R = right second conjunctival appendage, lateral branch; CAII(M)L = left second conjunctival appendage, medial branch; CAII(M)R = right second conjunctival appendage, lateral branch; CAIII = third conjunctival appendage; DCC = dorsal conducting canal; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; F = flange at base of crown of paramere; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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(Fig. 7c); evaporative areas occupying most New Guinea, or surrounding islands, with C. of metepisternum, and extending to excellens now known from as far north as Tai- mesepimeron (Figs 7c, d); ostiole moderate- wan (see below). The genus is also repre- ly-sized (Fig. 7d); peritreme elongate, most- sented in the Oriental region, with two ly rectilinear, with apex weakly curved at species known from Indonesia (Borneo and apex, gutter-like, medially sulcate (Fig. 7d). Sumatra). Coleotichus is also represented in Thoracic sterna: lateral margins strongly Pacific islands and archipelagos, distant from sulcate, prosternal and mesosternal keels Australia, including Samoa, Hawaii and the overlapping (Fig. 7c), with latter contiguous Marquesas. The most spectacular species of with explanate anterior margin of proepis- Coleotichus, C. blackburniae, is endemic to ternum, rounded (Fig. 7c); metasternal keels the Hawaiian Islands, where it breeds on thickened (Fig. 7c). Male Genitalia: py- koa, a native species of Acacia, where both gophore (Figs 7e, f, 8a) large, subquadrate to plant and insect are threatened, the jewel suboval, ventral margin convex; genital bug by green vegetable bug biological con- opening dorsal in orientation, broad, subo- trol agents (JOHNSON et al. 2005). val, lateral margins densely setose; para- Included species: meres (Fig. 8b), columnar stem, strongly C. adamsoni VAN DUZEE 1932 Marquesas hook-shaped crown, sometimes base of C. artensis (MONTROUZIER 1858) Australia, crown with flange; aedeagus (Figs 8c, d), New Caledonia, Fiji, asymmetrical (Fig. 8c, d) or symmetrical; Vanuatu, Samoa, Aru Island phallotheca squat (Fig. 8c), sclerotized, C. bakeri TAEUBER 1929 without processes; ejaculatory apparatus C. biroi SCHOUTEDEN 1905 New Guinea moderately developed (Fig. 8c); ventral C. blackburniae WHITE 1881 Hawaii conducting canal with 8-10 paired convolu- C. breddini SCHOUTEDEN 1905 Micronesia tions (Fig. 8c); ejaculatory reservoir short, C. bulowi SCHOUTEDEN 1905 Samoa oval (Fig. 8c); vesica short to elongate, ta- C. costatus (FABRICIUS 1787) Australia, pered towards apex, strongly arcuate; con- New Caledonia, Tonga junctival appendages (Figs 8c, d): CAI ab- C. excellens WALKER 1867 Australia, Fiji, sent; CAII asymmetrical (Figs 8c, d) or sym- New Guinea, New Caledonia, metrical, greatly enlarged, mostly membra- Samoa, Philippines, Taiwan, neous, distally bifurcate, CAII(L) with short Indonesia (Irian Jaya) (acute) or large (sickle-shaped) lobal scle- C. fuscus VOLLENHOVEN 1863 Ambon, rite, CAII(M) with short acute lobal scle- Ceram rite, or blunt bifid lobal sclerites; CAIII scle- C. marianensis USINGER 1946 Guam rotized (Fig. 8c, d) or membraneous, fused C. ornamentifer BERGROTH 1915 Borneo medially or separated, flange-shaped or S- C. sumatranus BREDDIN 1900 Sumatra shaped, sometimes bifid. Female Terminalia: ventrally oriented, co-planar; paratergites Remarks: Coleotichus is a distinctive VIII large, broad, subelliptoid, posterior genus of Elvisurinae, having greatly enlarged margin rounded; paratergites IX narrowly el- thoracic sternal keels, with the pro- and liptoid, medial margins truncate; gonocoxae mesothoracic keels overlapping (Fig. 7d). I transverse, undivided, posterior margin The genus shares some similarities with the concave, medial margins elevated. Sper- nominotypical genus Elvisura, with the matheca: short to moderately-sized proximal pronotum greatly expanded posteriorly, cov- fecundation canal; spermathecal reservoir ering the anterior margin of the scutellum narrowly dilated elliptoid, membraneous; (Figs 3c-e). The male genitalia resemble pump with dorsal and proximal flanges; bulb those of Elvisura, in the following ways: the oval. CAII have lateral and medial lobes with lob- Diversity and distribution: The highest al sclerites, and the CAIII are sclerotized and area of species diversity in Coleotichus is in fused medially (as in C. costatus and C. the Australian zoogeographic region, with artensis) (e.g., Figs 8c, d). LESTON (1953b) seven of the thirteen described species. A described the male genitalia of Elvisura irro- number of the species are widespread, with rata SPINOLA, which serve as a basis for this all three Australian species, found also in comparative hypothesis.

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WHITE (1839) described Coleotichus on (female genitalia); MCDONALD &CASSIS 1984: the basis of the Australian species, C. costa- 542 (synonymy) tus. SCHOUTEDEN (1904), in redescribing Coleotichus nigrovarius WALKER 1867: 2 (n.sp.); the genus, established three subgenera MCDONALD & CASSIS 1984: 542 (synonymy) (Coleotichus, Epicoleotichus and Para- Coleotichus testaceus WALKER 1867: 2 (n.sp.); STÅL 1873: 4 (synonymy) coleotichus) on the basis of the prosternal shape, declivity of the male pygophore, and Diagnosis: Coleotichus artensis is recog- the shape of the female genital plates. nised by the following combination of char- GROSS (1975) redescribed Coleotichus, and acters: AIV longest antennal segment; cal- used these subgeneric categories, giving cre- losite region of pronotum with four promi- dence to the shape of the prosternum in re- nent black spots; labium reaching apex of lation to the eyes. Our observations indicate metasternum; female abdominal SVII nar- that differences in these characters are more row caudally, ventral margin deeply con- indicative of species relationships, and not cave; males without abdominal sternal diagnostic at the genus level. glands; posterior margin of male pygophore excavate; CAII symmetrical, tripartite; MCDONALD &CASSIS (1984) added a CAII fused post-thecal margin; vesica elon- description of the male genitalia for gate; and, proximal sclerotization of sper- Coleotichus, and did not adhere to subgener- mathecal fecundation canal. ic categories, without any explicit explana- tion. Pending a more detailed comparative Description: Body elongate-ovoid (Fig. morphological investigation and phyloge- 3c); moderately sized, males 12.8-14.7 mm, netic analysis of Coleotichus, the sister- females 12.6-14.7 mm. species relationships are uncertain, and we Colouration. Body stramineous-brown know of no new evidence that supports the to dark-brown, sometimes with mottled ap- use of subgeneric categories. Aside from the pearance, punctures dark-brown to black, Australian species, the male and female sometimes with green iridescence (Fig. 3c). genitalia of the extralimital species of Head: stramineous-brown, sometimes darker Coleotichus have not been documented; a with clypeal margins fuscous, reaching pos- necessary prerequisite for any further classi- terior margin of vertex, posterior margin of ficatory hypotheses. head rarely transversely fuscous; anterior margin of pronotum sometimes dark brown Coleotichus artensis to fuscous; pronotal calli with prominent (MONTROUZIER 1858) (Figs 3c, 6) dark brown to fuscous spots; anterolateral Scutellera artensis MONTROUZIER 1858: 259 (n.sp.) angles of scutellum with black spot; scutel- Coleotichus artensis: STÅL 1873: 4 (new combina- lum often mottled, sometimes with medial tion); LETHIERRY &SEVERIN 1893: 15 (cata- darker brown oblique fasciae; thoracic pleu- logue); SCHOUTEDEN 1904: 6 (list; separate species); SCHOUTEDEN 1905: 336 (description); ra and abdominal sterna yellow-brown to SCHOUTEDEN 1907: 107 (Montrouzier type); dark-brown; deep punctures of proepisternal KIRKALDY 1909: 313 (catalogue); DISTANT 1920: keel sometimes with green iridescence; ab- 143 (New Caledonia); MCDONALD &CASSIS dominal spiracular-trichobothrial region 1984: 542, fig. 16 (description; spermatheca); contrastingly dark-brown; appendages con- CASSIS &GROSS 2002: 583 (catalogue; syn- colorous, pale to dark brown. onymy); SMITH 1978: 821 (defensive secretion) Coleotichus marginatus SIGNORET 1861: 59 (n.sp.); Texture. Dorsum with dense distribu- STÅL 1873: 4 (synonymy) tion of deep punctures, less so on head, more Coleotichus sordidus WALKER 1867: 1 (n.sp.); DIS- so on scutellum and pronotal disc, irregular- TANT 1899: 50 (as junior synonym of Coleotichus ly distributed; thoracic pleura with deep fuscus); CHINA 1930: 91 (Samoa); SMITH 1978: punctures submarginally on propleuron and 821 (defense); MCDONALD &CASSIS 1984: 542 mesopleuron. (synonymy) Coleotichus discrepans WALKER 1867: 2 (n.sp.); Vestiture. Dorsum near glabrous. An- MCDONALD 1961: 179, figs 24-28 (male geni- tennae: AI-AII(a) almost glabrous, AII(b)- talia); MCDONALD 1963a: 179, figs 24-28 (male AIV densely pilose, with short simple semi- genitalia); MCDONALD 1963b: 223, figs 15-16 erect setae. Legs: ventral margins of tibiae

310 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at densely pilose.Underside of body with scat- taceus from two specimens; the original descrip- tered short simple setae; lateral margins of tion listing them as: ‘a. New Caledonia. ‘From Mr abdominal sterna more densely pilose. Macgillvray’s collection’ and ‘b. Moreton Bay. From Mr Diggles’ collection’. The lectotype and Structure. Antennae: AI(a) little longer paralectotype specimens designated here, bear la- than AII(b), AIV longest segment. Labium: bels with these localities respectively. reaching apex of metasternum. Thoracic Other material examined: Queensland: 1Y, Ku- Pleura: external efferent system of metatho- randa, 5 December 1988, R Bejsak (AM); 1Y, racic glands extensive, peritreme elongate, Cairns, Crystal Cascades, 1 February 1989, J & M strongly recurved towards head. Abdominal Bugeja (AM); 1Y, Kuranda Range, State Forest, Venter: SIII with medial anteriorly project- 10 January 1967, DK McAlpine & G Holloway Y ing processes; posterior angles of SIV-SVII (AM); 1 , Bluewater Range, 16-v-1990, T = acute, particularly SVII; female abdominal Woodger (AM); 1 , 14 km ENE Heathlands, 11.41S 142.42E, 21 October 1993, P Zborowski SVII narrow caudally, posterior margin & DCF Rentz, at light, rainforest (ANIC); deeply concave; males without abdominal 2YY, Shiptons Flat, 15.47S 145.14E, 17 Octo- sternal glands. Male Genitalia: posterior ber 1980, T Weir (ANIC); 1Y, 9 km SSW Ku- margin of male pygophore excavate, with randa, 16.54S 145.37E, 25-26 November 1992, A medial setose patches; parameres with flange Calder & P Zborowski, at light (ANIC); 1Y, 2 at base of crown; phallotheca with pair of km N Kuranda, 16.48S 145.38E, 20 November subdistal thorn-like processes; ejaculatory 1981 (ANIC); 1Y, FP Dodd, 1907-54, ‘det. as reservoir heavily sclerotized; ductus seminis Coleotichus discrepans’ (BMNH); 1Y, Redlynch, distalis basally incrassate; CAI absent; CAII 21-30-viii-1938, RG Wind, ‘det. as Coleotichus symmetrical, tripartite mostly membraneous, discrepans’ (BMNH); : 4== Y CAII(L) with large, elongate sickle-shaped 1 , 3 to 5 km NE of Harrington, G Williams, lobal sclerite, CAII(M) with small, bifid lob- January 1989 & 1991, ex , lit- toral rainforest (AM); FIJI: 1=, Cuvie, ‘1920-82’, al sclerite, paired ventral CAII process with ii-1918, R Veitch, ‘det. as Coleotichus nigrovarius’ laterally oriented bill-shaped lobal sclerites; (BMNH); TONGA: ‘Distant Coll. 1911-383’, CAIII strongly sclerotised, medially fused, ‘det. as Coleotichus nigrovarius WALK.’ (BMNH); with apices arcuate, laterally directed, vesica VANUATU: 6== 6YY, Aneityum, 3 mi. NE elongate, extending beyond conjunctival ap- Anelgauhat, Red Crest, 200 ft, iii-1955, LE pendages. Female Terminalia: paratergites Cheesman, B.M. 1955217 (BMNH). VIII moderately sized, subtriangular, posteri- Distribution: Coleotichus artensis is or margin arcuate, medially separated; broadly distributed in the tropical regions of paratergites moderately sized, subtriangular, the Australian zoogeographic region; known medial margins truncate; gonocoxae I subtri- from Australia, Indonesia, Papua New angular. Spermatheca: proximal fecundation Guinea, Fiji, New Caledonia, Samoa and canal short, base sclerotized; bulb oval, heav- Vanuatu (CASSIS & GROSS 2002). Within ily sclerotized. Australia, it was known previously from the

Measurements. MCDONALD & CASSIS tropical parts of the and 1984: Table 1. Queensland. In this work, we greatly extend its range to the south, with new collections Type material examined: Coleotichus artensis from the mid-north coast of New South MONTROUZIER: Holotype, Y, ‘B.M. Hem. 365’, Wales (near Harrington) (Fig. 6). There is ‘New Caled’, ‘59-63’; Coleotichus discrepans also a significant distributional gap between WALKER: Holotype, Y, ‘B.M. Hem Type No. 366’, ‘Moreton Bay’ (BMNH); Coleotichus ni- the populations of the wet tropics and sub- grovarius WALKER: Holotype, Y, ‘B.M. Hem Type tropical regions of Queensland. No. 367’, ‘Fiji Isles, Ovalau’, ‘56-69’, ‘585’ Host plants and biology: Coleotichus (BMNH); Coleotichus sordidus WALKER: Holo- artensis was collected from the of the type, 1Y, ‘B.M. Hem Type No. 368’, ‘I of Pines’ littoral rainforest , Alphitonia excelsa (BMNH); Coleotichus testaceus WALKER: Lecto- type, Y, ‘New Caled’, ‘B.M. Hem. Type No. 365’, (Rhamnaceae), during two separate collec- ‘Coleotichus testaceus WALKER’s catal.’, ‘59-63’ tion events. This represents the first record- (BMNH; here designated); Paralectotype, Y, ed plant association for this jewel bug ‘Moreton Bay’, ‘artensis Montr.’ (BMNH; here species, but doubt remains as to whether designated). WALKER (1867) described C. tes- this is its food plant or breeding host.

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Remarks: MCDONALD & CASSIS (1984) Coleotichus costatus: WHITE 1842: 88 (new combi- redescribed C. artensis and gave a detailed nation); WALKER 1867: 1 (list); WALKER 1868: synonymy. It exhibits variability in colour, 505 (list); STÅL 1873: 4 (list); LETHIERRY & SEV- ranging from matt light brown to polished ERIN 1893: 15 (catalogue); SCHOUTEDEN 1904: 6 dark brown, otherwise it is homogeneous, (list); Kirkaldy 1909: 313 (catalogue; as junior synonym of Coleotichus unicolor)DISTANT 1920: particularly in the male genitalia. We re- 143 (New Caledonia); KUMAR 1965: 46 (male cently re-examined all the types, aside from genitalia); GROSS 1975: 81 (description); VAN C. marginatus, and found that a wider exam- DEN BERG 1980: 223-225 (biology; host plant); ination of extralimital specimens of this MCDONALD &CASSIS 1984: 9 (redescription; species is required to confirm the identity of synonymy; male genitalia: Figs 3-7, 9; female gen- this species and the above-mentioned syn- italia: Figs 7 & 8); CASSIS &GROSS 2002: 584 onymy. As the types are all females, doubts (catalogue) of the validity of the given species syn- Eurygaster costatus: VOLLENHOVEN 1863: 39 (new onymy prevail, as the male genitalia remain combination) the chief criterion for distinguishing species. Coleotichus unicolor DALLAS 1851: 5 (n.sp.); The holotype of C. sordidus, which was de- SCHOUTEDEN 1904: 6 (synonymy) scribed from a New Caledonian specimen Coleotichus pallidus VOLLENHOVEN 1863: 4 (n.sp.); SCHOUTEDEN 1904: 6 (synonymy) (Isle of Pines), is possibly deserving of species resurrection, as it is pale brown with- Diagnosis: Coleotichus costatus is recog- out patterned spotting, the pronotal humer- nised by the following combination of char- al angles are more rounded, and the postero- acters: AIII and AIV subequal, longest an- lateral angles of the abdominal venter are tennal segments; callosite region of prono- not pronounced, as in the other types. How- tum with pair of small submedial black spots ever, we have refrained from removing the (Fig. 3d); labium reaching anterior margin latter from synonymy pending a generic re- of metacoxae; female abdominal SVII broad vision of Coleotichus, and found that its pale caudally, ventral margin shallowly concave; colouration is consistent with individuals males with abdominal sternal glands on found in Australian populations that we ob- SIV-VI (Figs 2a-c); posterior margin of py- served. gophore arcuate; CAII asymmetrical (Figs Coleotichus artensis most resembles C. 8c, d); CAII fused post-thecal margin (Figs costatus in colour and size, although the 8c, d); vesica elongate (Figs 8c, d); and, mean lengths of both sexes of C. artensis are basal sclerotisation of spermathecal fecun- smaller and the maximal width of males is a dation canal. little larger. The male genitalia of C. artensis Description: Body elongate-ovoid (Fig. are discrete with the CAII tripartite; a char- 3d); moderately large, males 12.6-15.5 mm, acter state which does not occur in all other females 13.4-17.2 mm. species of Coleotichus investigated. Like oth- Colouration. Body yellow-brown to or- er species of Coleotichus, the male genitalia ange-brown, sometimes with dusty appear- of C. artensis exhibit little intraspecific vari- ance, often with darker brown patterning on ation. Its closest relationships may lie with scutellum (Fig. 3d), sometimes with red extralimital species such as C. biroi, which it highlighting, jugal margins, anterolateral resembles on the basis of external characters margins of pronotum and embolium most and size. The female genitalia of C. artensis often contrastingly paler yellow. Head: uni- and C. costatus are alike, with the proximal formly yellow-brown, punctures either fus- sclerotization of the fecundation canal, a fea- cous or with green iridescence; underside of ture not found in C. excellens. head yellow; antennae, AI yellow, AII-AIV Coleotichus costatus (FABRICIUS 1787) yellow-brown to orange-brown; labium yel- (Figs 3d, 6, 7, 8) low to yellow-brown distally, with stylets black. Pronotum: pale red fasciae submar- Cimex costatus FABRICIUS 1787: 282 (n.sp.); 1805: pl. 3 fig. 5 (habitus); GERMAR ginal to anterolateral margins; callosite re- 1839: 74 (list) gion sometimes with pair of small submedial Tetyra costatus: FABRICIUS 1803: 135 (new combi- spots. Hemelytra: exocorium and base of nation) clavus with dark punctures, often with green

312 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at iridescence, exocorium sometimes with pale land’ [label data as ‘New Holl’], ‘B.M. Hem. Type red longitudinal fascia. Scutellum: postero- No. 364’, ‘unicolor DALLAS’ (BMNH). lateral margins with fuscous punctures, of- Other material examined: Queensland: 2YY, ten with green iridescence. Legs: uniformly 146 km NW Quilpie. 25:858S 143.399E, 230m, yellow-brown to orange-brown. Thoracic G Cassis, RT Schuh & R Silveira, 3 November 1998, ex Acacia stowardii MAIDEN, Site 98-20 pleura and abdominal sterna: uniformly yel- (AM); 3== 2YY, 73.7 km E Betoola, 25.591S low-brown, often with red spotting. 141.399E, 180m, G Cassis, RT Schuh & R Sil- Texture. Body densely punctate, punc- veira, 3 November 1998, at light. (AM); New tures shallow; mostly evenly distributed, lin- South Wales: 1=, Kinchega National Park, G Y early arranged on head, absent from cal- Cassis, 28 April 1995, ex grass (AM); 1 , Kinchega National Park, Cawndilla Camp- losite region and anterolateral margins of ground, 32°33’S 142°12’E, 100m, G Cassis & RT Pronotum. Schuh, 28 October 1995, Site 95-34, at light. Vestiture. Dorsum glabrous; underside (AM); 2== 4YY, Moppin-Aveymore Road, of body almost glabrous, with scattered dis- 28°53’26“S 149°51’30“E, 400 m S of junction at tribution of short, simple setae on abdomi- Dolgelly Bore, R Harris & T Moulds, ex Acacia nal venter, more so caudally. Antennae: AI- pendula (AM); 15 km NE by N Moree, 29°20’S 149°56’E, 21 April 1981 (ANIC); Australian AII near glabrous; AIII-AIV with moderate Capital Territory: 52 specimens, Griffith, 17 distribution of short simple semierect setae. February 1969, TG Campbell & CJ Shepherd, Legs: sparse distribution of simple short (ANIC); Southern Australia: 1Y, Cadelga semierect setae. Homestead, 26.089S 140.410E, 150m, G Cassis, Structure. Antennae: AIII and AIV RT Schuh & Silveira, 4 November 1998, Site 98- 24, at light (AM); 2== 2YY, Mt Serle district subequal, longest antennal segments. Labi- (near Gammon Ranges National Park), um: reaching anterior margin of metacoxae. 30°33’15“S 138°50’13“E, 567 m, 8 November Abdominal Venter: males with sternal 2001, RT Schuh, G Cassis, & M Schwartz, ex glands on abdominal SIV-VI (Figs 2a-c). Acacia victoriae (AM); Western Australia: 1=, Male Genitalia: pygophore lozenge shape NW Coastal Highway, 36km N Kalbarri Road, (Figs 7e, f, 8a); ventral margin of pygophore 27.616°S 114.683°E, 500m, G Cassis & RT arcuate (Fig. 7e); parameres with strongly Schuh, 28 October 1996, Site 96-38, at light hooked crown, with flange at base of crown (AM); 1Y, 20 km S of Menzies, G Cassis & RT (Fig. 8b); CAI absent (Fig. 8c); paired CAII Schuh, 28 October 1996 (AM); 3== 6YY, asymmetrical (Fig. 8c, d), greatly enlarged, Moorine Rocks, 11.7 km N Great Eastern High- mostly membraneous, distally bifurcate, way on Noongar Rd, 31.228°S 118.986°E, 345 m, RT Schuh, G Cassis, H Brailovsky & A Asquith, right CAII(L) with enlarged, sickle-shaped 4 December 1997, ex Acacia saligna (LABILL.) lobal sclerite, right CAII(M) with blunt bi- H.L. Wendl, Site 97-01 (AM); 21== 33YY, fid lobal sclerite, left CAII(L) with moder- 55.6 km SE Southern Cross, 31.589°S 119.592°E, ately-sized, arcuate lobal sclerite, left 470m, RT Schuh, G Cassis, H Brailovsky & A CAII(M) bifid with short subdistal acute Asquith, 4 December 1997, ex Acacia consan- lobal sclerite and small apical serrate lobal guinea R.S. Cowan & Maslin, Site 97-03 (AM); sclerite; CAIII small (Fig. 8c, d), flange- 1Y, District, Shay Gap Rd, 15.1 km NE shaped, fused, heavily sclerotized, vesica ex- Muccan Homestead, 20.222S 120.149E, 130m, G tending beyond conjunctival appendages. Cassis & R Silveira, 27 May 1999, at light. (AM); Female Venter: posterior margin of SVII bis- 1Y, adjacent to N boundary of Lake Shaster Na- inuate, medially thickened. Female Termi- ture Reserve, 33.833°S 120.916°E, 40m, RT Schuh, G Cassis & R Silveira, 27 November nalia: paratergites VIII large, medial mar- 1999, ex , Site 99-42 (AM); 2 gins broadly contiguous. Spermatheca: fe- larvae, Rossiter Bay, Cape Le Grande National cundation canal short, proximally incrassate Park, 3 m, 33°58.0345’S 122°16.0457’E, 23 No- and heavily sclerotized; and, spermathecal vember 1999, RT Schuh, G Cassis & R Silveira, bulb oval, heavily sclerotized. Site SWA99-29, ex Acacia cyclops (AM); 1Y, Walsh Point, 14.34 125.51E, 17 May 1983, I Measurements. MCDONALD & CASSIS Naumann & JC Cardale (ANIC); 5 specimens, 1984: Table 1. Red Bluff, 28 November 1971, N McFarland, ex Type material examined: Coleotichus unicolor green pods of Acacia sp. (ANIC); 3 specimens, DALLAS: Holotype, =, ‘North Coast of New Hol- Mt Magnet, 6 December 1978, K&E Carnaby

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(ANIC); 2 specimens, 1 km W Jimberiana Hill, more generally within the Heteroptera. Norseman, 32°09’S 121°48’E, 11 January 1993, None-the-less the affinities of C. costatus ED Edwards & ES Nielsen (ANIC). are more with C. artensis, which possess Distribution: Coleotichus costatus occurs fused CAIII (cf. C. excellens; paired, segre- in arid and semi-arid regions of Australia, gated CAIII). In addition, the two former primarily in temperate Australia (Fig. 6). It species are similar externally, with the body is also known from tropical Australia, in- smaller, the ventral margin of the pygophore cluding the Pilbara district of Western Aus- emarginate, and the base of the spermathe- tralia and the wet tropics of Queensland. cal fecundation canal heavily sclerotized. MCDONALD & CASSIS (1984) record it from the Northern Territory, although no precise Coleotichus excellens (WALKER 1867) localities are currently known. It has also (Figs 3e, 6)

been recorded from New Caledonia and Coleotichus excellens WALKER 1867: 3 (n. sp.); Tonga (CASSIS & GROSS 2002). LETHIERRY & SEVERIN 1893: 15 (catalogue); DIS- TANT 1899: 30 (Australia, Samoa); HUTTON Host plant records and biology: 1904: 332 (erroneous New Zealand record); Coleotichus costatus is known to feed on sev- SCHOUTEDEN 1904: 6 (list); SCHOUTEDEN 1905: en species of Acacia (consanguinea, cyclops, 328 (description); BERGROTH 1908: 139 (descrip- ligulata pendula saligna stowardii victori- , , , and tion); KIRKALDY 1909: 313 (catalogue); MCDON- ae; Table 1) in Australia, primarily in semi- ALD &CASSIS 1984: 540, Figs 11-15 (description; arid and arid regions. It can be abundant female genitalia); CASSIS &GROSS 2002: 585 when the seed set of Acacia species is high (catalogue) (e.g. Southern Cross locality). It appears to Coleotichus borealis DISTANT 1899: 31 (n.sp.) be opportunistic and probably feeds on addi- New Synonymy tional Acacia species, depending on seed Coleotichus handlirschi SCHOUTEDEN 1905: 332 (n. availability. WHITNEY & STANTON (2004) sp.); MCDONALD &CASSIS 1984: 540 (syn- report that C. costatus primarily feeds on onymy) Coleotichus (Epicoloeotichus) schultzei TAEUBER Acacia ligulata in the Kinchega National 1929: 221 (n.sp.) New Synonymy Park in western New South Wales, but oc- casionally feeds on the seeds or fruits of Diagnosis: Coleotichus excellens is recog- Dodonaea viscosa (Whitney pers. comm.). nised by the following characters: AIII and The Lambertia inermis plant association AIV subequal, longest antennal segments; (Table 1) is probably a sitting record. As labium reaching abdominal sterna IV, LII- with other species of jewel bugs, GROSS IV subequal in length, longest segments; (1975) reports that C. costatus can be found callosite region of pronotum with pair of on the ground, but only occasionally on small submedial black spots; scutellum with vegetation. Our observations do not support antero-sublateral orange calli; scutellum of- this contention, with C. costatus commonly ten with six small black spots; female ab- encountered feeding on pre-dispersed seeds, dominal SVII broad caudally, ventral mar- upon plants. gin shallowly concave; posterior margin of male pygophore convex; males without ab- Remarks: SCHOUTEDEN (1904) estab- dominal sternal glands; CAII symmetrical, lished the synonymy for Coleotichus costatus, bipartite; CAII separated; and, vesica short. which has been supported by all subsequent authors. GROSS (1975) redescribed this Description: Body elongate-ovoid, ta- species including a habitus illustration. MC- pered towards terminalia; large species, males DONALD &CASSIS (1984), also redescribed 17.67-21.50 mm, females 18.17-21.83 mm; C. costatus, documenting the male and fe- Colouration. Body uniformly yellow- male genitalia for the first time. CASSIS & brown (Fig. 3e), with yellow midline, often GROSS (2002) gave a comprehensive syn- with green or purple iridescent punctures. onymy for the species. Head: punctures of jugal margins strongly Coleotichus costatus is best defined by the iridescent green. Pronotum: callosite region distinctive aedeagus, with the CAII asym- with pair of submedial black spots. Scutel- metrical (Figs 8c, d). This condition appears lum with pair of antero-sublateral black to be unique in the Scutelleridae, and even spots, and medial black spot.

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Texture. Dorsum uniformly punctate; males, and the specimen labeled as ‘Coleotichus with moderately deep punctures; anterolat- excellens WALKER (type)’ and ‘B.M. Hem. Type eral margins of pronotum broadly impunc- No. 369” is designated as the lectotype; the other tate; anterolateral calli of scutellum smooth; two specimens as paralectotypes; Coleotichus bore- alis DISTANT: Holotype, Y, ‘’, ventral aspect of jugae and antennifers ‘Coleotichus borealis DIST. (type)’, ‘B.M. Hem. deeply punctate; thoracic pleura moderately Type No. 370’ (BMNH); Paratype: =, same data punctate. as holotype, ‘Coleotichus borealis DIST.’ (BMNH); Vestiture. Dorsum glabrous. Antennae: Coleotichus schultzei TAEUBER: Holotype, =, ‘Suri- AI-AII(a,b) near glabrous, AIII-AIV with gao’, ‘Minando’, ‘13351’, ‘Taeuber Coll. 1949- 474’ ‘Holotype Coleotichus (Epicoleotichus) moderate distribution of short setae. Legs: schultzei TAEUBER 1929’, ‘B.M. Hem. Type No. femora with sparse distribution of short se- 602’ (BMNH; genitalia dissected, in microvial). tae; tibiae with moderate distribution of Other material examined: Queensland: 1=, short setae, more so on ventral surface. Ab- Black River, 20 km N Townsville, 14-i-1990, T dominal Venter: scattered short simple Woodger, at light (AM); 1=, Gap Creek, 15°51’S semierect setae, more so on terminalia. 145°20’E, 29 May 1994, P Zborowski, ex light, rainforest (ANIC); 1=, 1 km E Mt Cook, 13°30’S Structure. Antennae: AI and AII(a), 145°16’E, 13 October 1980, T Weir (ANIC); and AIII and AIV subequal in length. Labi- 4==, Warraber Island, 10°12’S 142°49’E, March um: elongate, reaching abdominal SIII, LII- 1978 (ANIC); 1=, Mt Webb National Park, IV roughly subequal in length. Pregenital 15°04’S 145°07’E, 27-30 April 1981, A Calder Abdomen: SIII medially sulcate, contiguous (ANIC); 2YY, 12 km SE Daintree, 16°19’S with thoracic sternal keel. Male Genitalia: 145°24’E, 22 November 1981, J Balderson (AN- Y ventral margin of pygophore arcuate; genital IC); 1 , 32 km N by W of Isabella Creek, 15°18’S 145°00’E, 230 m, 22 May 1975, IFB opening broad; parameres with hooked Common & ED Edwards (ANIC); 1Y, Mackay, crown, tip blunt, with flange at base of AJ Turner, 1905-125 (BMNH; identified as crown; CAI absent; CAII symmetrical, Coleotichus borealis; Northern Territory: 1Y, mostly membraneous, bifid, CAI(L) and Smith Point, Cobourg Peninsula, 11°07’S CA(M) with acute lobal sclerites; CAIII, 132°08’E, 3-21 February 1972, RC Lewis, ex wa- heavily sclerotized, S-shaped, segregated. tertrap (ANIC); 1= 1Y, 1 km N Cahills Cross- Female Terminalia: ventrally oriented, co- ing, E Alligator River, 12°25’S 132°58’E, 31 Oc- planar; paratergites VIII large, posterior tober 1972, M Upton & Barrett (ANIC); New Caledonia: Province Sud, 1 km S Poya, margin arcuate, medially contiguous; 21°21’19.4“S 165°10’8.1E, 61 m, 27 April 2005, paratergites IX large, subelliptoid, medial G Cassis, MA Wall, N Tatarnic & GB Monteith, margins arcuate; gonocoxae I large, subtri- ex Ficus sp. (AM); Papua New Guinea: 1=, Port angular, posterior margin weakly convex. Moresby, Broko, Cent[ral Dist[trict], 15-iii-1959, Spermatheca: fecundation canal elongate; ex Hibiscus rosasinesis, JJH Szent-Ivany (BMNH); spermathecal reservoir large, oval; bulb 1=, Port Moresby, 26 February 1969, R Lossin round, moderately sclerotized. (AM); 1Y, Amazon Bay area, Doveta, 2400 ft, 24.vii-11.ix, 1962, WW Brandt (ANIC); 1=, Measurements. MCDONALD & CASSIS Kokoda, 1,200 ft, ix-1933, LE Cheesman, B.M. 1984: Table 1. 1934-321’ (BMNH); 2==, Madew, St Joseph’s River, 2000-3000 ft, W Stalker, 1909-22 Type material examined: Coleotichus excellens (BMNH); Solomon Islands: 1Y, Malaita, Auki WALKER: Lectotype, Y, ‘124’, ’56-69’, ‘B.M. Harbour, 24-xii-1965, Roy. Soc. Exped. Brit. Hem. No. 369’, ‘Coleotichus excellens WALKER Mus. 1966-1’, ‘169’, ‘Coleotichus excellens WALK- (type)’ (BMNH; here designated); Paralecto- ER’s catal.’ (BMNH; identified as Coleotichus han- Y types, 1 : ‘56-69’, ‘Coleotichus excellens WALK- dlirschi); 4YY, Guadalcanal, PJM Greenslade, ER’S catal.’, ‘Coleotichus excellens: Det. J. Cassis’ xii-1964, 3-I-1965, 8-I-1965 (BMNH); Fiji: Y (BMNH; here designated); 1 , ‘55-69’, ‘127’, 3YY, Rotuma Is., 17-27-iv-1971, GS Robinson, ‘Coleotichus excellens WALKER’S catal.’ (BMNH; B.M. 1972-46 (BMNH); 1Y, Suva, 20-I-1030, here designated). WALKER (1867) listed three B.M. 1948-548 (BMNH); Philippines: 2== specimens in his original description, as follows: 2YY, Balabae, Dalwan Bay, Noona Dan Exp. ‘a,b. ___ ? From Mr Macgillvray’s collection’ and 61-62, 18-30 to 23-30, [1962], ex mercury light c.___ ? Presented by Sir John Liddell’. Neither (BMNH; identified as Coleotichus borealis); 1Y, the original description, nor the label data indi- C.S. Banks 1908-228, Acc. No. 6377, Bu. of Sci., cate a type locality. All the specimens are fe- P.I. (BMNH; identified as Coleotichus borealis);

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Indonesia: 1=, [Irian Jaya], Waigeu Camp Nok, been examined. A dissected male specimen N Dutch New Guinea, 2,500 ft, LE Cheesman, of C. bulowi in the BMNH, which is similar B.M. 1938-593 (BMNH). in colouration and size to C. blackburniae, Distribution: Coleotichus excellens is and its aedeagus is very similar to that of C. broadly distributed in the Australian zoo- excellens (CAII bifurcate, branches near geographic region, extending to the north- symmetrical, with conical lobal sclerites). east of Australia (Solomon Islands, New Caledonia, Fiji, and Samoa), and to the Ori- Solenotichus MARTIN 1897 ental region, including the Philippines and (Figs 3b, 6, 9, 10)

Taiwan. We provide the first record of this Solenotichus MARTIN 1897: 264 (nov. gen.); DIS- species from Indonesia, Papua New Guinea TANT 1899: 31 (description); KIRKALDY 1904: 280 and Fiji. Within Australia, its found prima- (note); SCHOUTEDEN 1904: 7 (description); rily in the tropical regions of the Northern GROSS 1975: 86 (description); MCDONALD & Territory and Queensland, extending in the CASSIS 1984: 545 (description); CASSIS &GROSS latter as far south as Rockhampton (Fig. 6). 2002: 585 (catalogue) CASSIS & GROSS (2002) reported this Damelia DISTANT 1899: 31 (gen. nov.; junior species from New Zealand, in an erroneous homonym of Damelia CLARK 1864 [Coleoptera] Damellera KIRKALDY 1904: 280 (nom. nov. for reference to HUTTON (1904). LARIVIÈRE & Damelia DISTANT 1899); KIRKALDY 1909: 313 LAROCHELLE (2004) did not mention it in (synonymy) their New Zealand catalogue. Type species: Solenotichus: Solenotichus brevipes Host plants and biology: There are no MARTIN 1897, monotypy; Damelia: Sphaerocoris host plants recorded in the literature for circuliferus WALKER 1867, monotypy Coleotichus excellens. We recently collected a Diagnosis: Solenotichus is recognised by large series from an undetermined Ficus the following combination of characters: species in New Caledonia, at a roadside rest- body ovoid (Fig. 3b); dorsum strongly con- stop, where there were few shrubs or trees. vex; posterior margin of pronotum arcuate We also record specimen from New Guinea (Fig. 3b); metathoracic peritreme narrow, (see above) which was collected on an or- moderately elongate, medially sulcate, namental Hibiscus. strongly raised distally (Fig. 9d); male ab- Remarks: New synonymy for this species dominal sternal glands submedially on SIV- is proposed, with Coleotichus borealis DIS- SVII; ventral conducting canal of ejaculato- TANT and Coleotichus schultzei, established as ry apparatus almost straight (Figs 10c, d); junior synonyms. The external features of ejaculatory reservoir small, circular (Fig. the holotypes are identical to those of the 10c); CAI absent; CAII bifurcate, with tri- holotype of C. excellens. The male genitalia angular lobal sclerites; CAIII sclerotized, of the holotype of C. schultzei, and a para- separate, apices laterally recurved (Figs 10c, type of C. borealis were examined, and are d); vesica elongate, arcuate (Figs 10c, d); consistent with those found in C. excellens. spermathecal fecundation canal short; and, spermathecal reservoir oval, wrinkled. MCDONALD & CASSIS (1984) gave the most recent redescription of Coleotichus ex- Description: Body small to moderately cellens, including the description and illus- sized; ovoid (Fig. 3a); dorsum strongly con- tration of the male and female genitalia. It is vex; moderately convex ventrally; orange distinct from the other Australian species of to orange-brown, with patterned markings; Coleotichus, and can be distinguished from dorsum moderately to heavily punctate, less them by the larger body, dorsal markings, so ventrally. Head: subhemispherical, elongate labium (with modified abdominal strongly transverse (Fig. 9a); moderately SIII housing), and the segregated CAIII. Its deflexed (Fig. 9b); lateral margins of jugae affinities are unknown, as it differs from oth- convex, subcarinate (Fig. 9b). Pronotum: er Coleotichus species in that the ventral transverse, subtrapezoidal, strongly convex conjunctival appendages are not fused. This (Fig. 3b); anterior margin concave; antero- species approaches the Hawaiian species, lateral margins strongly divergent, weakly , in size, however, convex, rounded in profile; posterolateral the male genitalia of this species have not margins weakly convex; posterior margin

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Fig. 9: Scanning electron micrographs of key characters of Solenotichus circuliferus (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ea = evaporative area; Ms(k) = mesosternal keel; Pe = peritreme; P(r) = right paramere. Scale bars = 1 mm.

arcuate. Scutellum: broad, covering most of SIV-SVII, unpaired on SVII. Male Geni- forewings and abdomen, U-shaped, strongly talia: ventral margin of pygophore truncate convex (Fig. 3b); heavily punctate; anterior (Fig. 9e); genital opening large, transverse, region raised, posteriorly strongly declivent. moderately pilose laterally (Figs 9f, 10a); Thoracic sterna: moderately elevated, parameres, large, stem broad, crown with thickened keels, not overlapping (Fig. 9c); large hook, apex blunt (Fig. 10b); aedeagus, anterior margin of proepisternum explanate ejaculatory apparatus relatively simple and (Fig. 9c). Thoracic pleura: external efferent moderately short; ventral conducting canal system of metathoracic glands well-devel- almost straight (Figs 10c, d); ejaculatory oped (Fig. 9c, d); ostiole large; peritreme reservoir small, circular (Fig. 10c); ductus moderately elongate, narrow, medially sul- seminis distalis without basal tumescence; cate, strongly raised distally; evaporative ar- CAI absent; CAII symmetrical, mostly eas extending beyond peritreme and reach- membraneous, distally bifid, CAII(L) and ing mesepimeron (Figs 9c,d). Pregenital CAII(M) subequal in size and shape, both Abdomen: abdominal sternal glands on with large triangular, lobal sclerites (Figs

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Fig. 10: Male genitalia of Solenotichus circuliferus (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII = third conjunctival appendage; DCC = dorsal conducting canal; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; F = flange at base of crown of paramere; GO = genital opening; S = stem of paramere; Sg = secondary gonopore; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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10c, d); CAIII prominent, segregated, scription); KIRKALDY 1909: 313 (catalogue); heavily sclerotized, arcuate, apices laterally GROSS 1975: 86 (description); CASSIS & GROSS recurved (Figs 10c, d); vesica elongate, re- 2002: 586 (catalogue) curved (Figs 10c, d). Female Terminalia: Diagnosis: Solenotichus circuliferus is paratergites VIII subtriangular, medially ta- recognised by the following combination of pered; paratergites IX large, subtriangular, characters: body mostly orange-brown, with medial margins rounded; gonocoxae I large, brown markings mediolaterally on scutellum subrectangular, posterior margin concave. (Fig. 3b); endocorium dark; AI-AIII sube- Spermatheca: proximal fecundation canal qual in length; LI and LIV subequal, LII short; spermathecal reservoir oval, with longest segment; male and female genitalia wrinkled texture; distal fecundation canal, as in generic description (Figs 10a-d). with proximal flange only; bulb oval, mod- Description: Body ovoid; moderately- erately sclerotised. sized species, males 9.3-10.7 mm, females Diversity and distribution: Solenotichus 8.7-11.8 mm. is a monotypic genus, endemic to Australia. Colouration. Body yellow-brown to red- Included species: brown, often with lateral pair of arcuate S. circuliferus (WALKER 1867) Australia darker markings at half length of scutellum, sometimes with yellow tinge anterior to Remarks: Solenotichus is a distinctive dark markings; punctures often red; ap- genus of Elvisurinae, due to its oval and pendages concolorous with body, sometimes highly convex body, elongate and arcuate with tibiae more red; exocorium sometimes vesica, separated CAIII, simple ejaculatory darker at base, endocorium mostly fuscous. apparatus (without convoluted ventral con- ducting canal) (Figs 10c, d), and moderate- Texture. Body with dense distribution ly elongate, raised metathoracic peritreme of shallow punctures. (Figs 9c, d). The separate CAIII of Soleno- Vestiture. Body almost glabrous; ap- tichus also occur in Coleotichus excellens, but pendages sparsely setose. differ significantly in shape (arcuate versus Structure. Antennae: segments roughly S-shaped). On this character system alone, subequal in length, AIII and AIV a little the monophyly of Coleotichus is brought in- longer than previous segments. Labium: to question, however the truncate posterior reaching apices of metacoxae, LII longest margin of the pronotum, and other aspects segment; and, male (Figs 10a-d) and female of the male aedeagus, indicate other possible genitalia as in generic description. affinities. In summary, the sister-taxon rela- tionship of Solenotichus is not established on Measurements: MCDONALD &CASSIS the basis of the characters examined. 1984: Table 2. = GROSS (1975) and MCDONALD &CAS- Type material examined: Lectotype, , Australia, SIS (1984) redescribed the type species, and ‘58-124’, ‘B.M. Hem. Type No. 372’ (BMNH; = confirmed its synonymy, but did not com- here designated); Paralectotype, , same data as lectotype (BMNH; here designated). WALKER ment on its relationships. CASSIS & GROSS (1867) described from two male specimens, pre- (2002) provided a comprehensive synonymy sumably from the same locality (given as ‘Aus- for the genus. tralia’ in original description and label data). Other material examined: Queensland: 1=, Bluff Solenotichus circuliferus Range, via Biggenden, 1000 m, 16 August 1974, (WALKER 1867) (Figs 3b, 6, 9, 10) H Frauca (ANIC); 1=, Bluff Range, via Biggen- Sphaerocoris circuliferus WALKER 1867: 7 (n.sp.) den, 16 December 1971, H Frauca (ANIC); 1Y, Solenotichus brevipes MARTIN 1897: 264 (n.sp.); Coast Range, via Boompa, 18 April 1977, H MCDONALD & CASSIS 1984: 545, Figs 24-29 Frauca (ANIC); Southern Australia: 1Y, 51 km (synonymy; male and female genitalia) NW of Morgan, 33.835S, 140.800E, 150m, G Damelia circuliferus: DISTANT 1899: 32 (new com- Cassis, RT Schuh & G Gross, 1 November, 1995, bination) Site L95-44, at light (AM); Northern Territory: Solenotichus circuliferus: SCHOUTEDEN 1904: 7 2== 2YY, Alice Springs, 5 September 1992, R (new combination); BERGROTH 1908: 139 (de- Patterson, at light (ANIC).

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Distribution: Solenotichus circuliferus is neous, and at times with paler stripes or broadly distributed across Australia, includ- spots, sometimes mottled, rarely black, oc- ing Queensland, New South Wales, Victo- casionally with patterning on scutellum ria, South Australia and the Northern Terri- (e.g., Fig. 11a-c); larvae dull in colouration; tory (Fig. 6). It is known as far north as pro- and mesothoracic sterna flat to weakly Rockhampton in Queensland and Alice elevated, never keel-like, metasternum Springs in central Australia. mostly flat, rarely a little raised; anterior margin of proepisternum weakly to moder- Host plants and biology: This species is ately explanate; pronotum often subtrape- relatively rare in collections and its biology zoidal, with anterolateral margins ex- remains unknown. Based on specimen local- planate, and anterior angles spinose, some- ity data, CASSIS & GROSS (2002) recorded it times more rounded, with lateral margins from the Desert Gum, Eucalyptus gongylo- evenly arcuate, rarely, with a medial notch, carpa (Myrtaceae). There is no evidence with anterior angles anteriorly projected, in whether this record is a food-preference, front of posterior margin of eyes; external ef- overwintering site or sitting record. ferent system of metathoracic glands often Remarks: GROSS (1975) and MCDON- greatly reduced, without peritreme, with ALD & CASSIS (1984) redescribed Soleno- prominent rounded ostiole, laterally orient- tichus circuliferus, with the latter authors ed, evaporative areas often greatly reduced documenting the male and female genitalia. to absent, sometimes external efferent sys- A female from Western Australia has not tem more well developed, with short oval been included in this work. This specimen is peritreme, elevated, bounded by moderate significantly larger than all other specimens distribution of evaporative areas, also occur- of S. circuliferus, but the other morphologi- ring on mesepimeron; sometimes males ab- cal attributes are consistent with the defini- dominal androconial glands present (e.g., tion of the species. Figs 2a-c, 12e); pregenital abdomen without stridulatory vittae; ventral surface of py- Subfamily Odontotarsinae gophore caudally (e.g., Fig. 12e) or ventral- Scutelleroides, Odontoscelidae DALLAS 1851: 54 ly oriented; parameres hook-shaped, often (new family) weakly (e.g., Fig. 14b); ejaculatory apparatus Pentatomidae, Scutellerina, Odontotarsaria STÅL weakly to moderately-developed, usually 1872: 32 (new suprageneric taxon) without prominent ventral conducting Pentatomidae, Scutellerinae, Odontotarsaria: canal (not convoluted), ejaculatory reser- SCHOUTEDEN 1904: 32 (synonymy of Eurygas- voir usually small and oval (e.g., Fig. 14c); traria 1872 STÅL and Odontoscelaria STÅL 1872) CAI present or absent (e.g., Fig. 14c); CAII Pentatomidae, Scutellerinae, Odontotarsini: unbranched or bifurcate (e.g., Figs 14c, d), KIRKALDY 1909: 263 (catalogue) either sclerotized or membraneous with lob- Pentatomidae, Scutellerinae, Eurygastrini, Odon- al sclerites; CAIII medially contiguous (Fig. totarsaria + Odontoscelaria: LESTON 1952d: 13 (new suprageneric classification) 11d); spermathecal fecundation canal short; Pentatomidae, Odontoscelinae: MCDONALD and, spermathecal reservoir oval to weakly 1966: 67, 68 (new suprageneric classification) dilated. Scutellerinae, Odontotarsinae: MCDONALD & Remarks: STÅL (1872) described the CASSIS 1984: 547 (Australia) Odontotarsinae (as Odontotarsaria) a tribe Diagnosis: The Odontotarsinae are of the scutellerids, and redefined another recognised by the following combination tribe, as the Odontoscelaria, for a group of characters: small (e.g., Figs 11a-c) to moder- genera mostly from the Eastern Hemisphere. ately-sized species; ovoid (e.g., Figs 11a-c), He separated these taxa primarily on the elongate-ovoid or elongate, rarely strongly shape of the head (elongate versus semi- tapered at terminalia; body strongly punc- oval), the lateral margins of the pronotum, tate; often with setigerous punctures, and and degree of carination of the metathoracic sometimes with sericeous setae (Figs 13a-i), sterna. SCHOUTEDEN (1904) synonymised rarely with body extremely setose with elon- these two taxa and the Eurygastraria sensu gate setae; mostly grey-brown to dark brown STÅL, and recognised the Odontotarsaria as species (e.g., Figs 11a-c), sometimes strami- the valid name, even though the Eurygastri-

320 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at dae DALLAS and Odontoscelidae DALLAS potentially includes the Australian genus have priority. He recognised the tribe on the Morbora (see below for discussion). Another basis of absence characters, such as the ab- putative clade, is represented by a group in- sence of stridulatory vittae on the pregenital cluding and potentially related to Odonto- abdomen, the lack of a keel-like carination tarsus (e.g, Phimodera, Xerobia), which lack on the thoracic sterna, as well as characters the male sternal glands, and sometimes have of general application, such as the biconvex excavate anterolateral margins of the body. KIRKALDY (1909) followed SCHOUTE- Pronotum. These ideas are reminiscent of DEN (1904), and grouped the three above- STÅL’S Odontotarsaria and Odontoscelaria, mentioned suprageneric taxa within the sin- and require explicit character testing. We gle tribe, and included the same taxa within retain the use of the Odontotarsinae in the it. LESTON (1952d) reverted to the Stålian broad sense in this work, acknowledging the classification in part, by distinguishing these above limitations. three groups as subtribes, within the tribe Eurygastrini. He hypothesised that these Morbora DISTANT 1899 taxa were clearly differentiated from both (Figs 2d, e, 11, 12, 13, 14, Table 5) his Scutellerini and Pachycorini on the ba- Morbora DISTANT 1899: 47 (gen. nov.); DISTANT sis of the male genitalia, suggesting that the 1899: 47 (description); BERGROTH 1904: 355 Scutellerini-type was distinctive, and more (note); SCHOUTEDEN 1904: 86 (description); derived from the pentatomid-type, the latter KIRKALDY 1909: 265 (catalogue); GROSS 1975: 88 being more characteristic of pachycorines (description); MCDONALD &CASSIS 1984: 538, 547 (key; revision); CASSIS &GROSS 2002: 586 and eurygastrines. (catalogue) Modern Australian workers (e.g., GROSS Type species: Morbora australis DISTANT 1899 1975; MCDONALD & CASSIS 1984) have re- monotypy jected this approach, mainly based on MC- Diagnosis: Morbora is recognised by the DONALD’s (1966) work on North American following characters: body oval; head semi- taxa, who regarded the eurygastrines as dis- circular, with dentate margins (Figs 11a-c, tinct from odontotarsines. SCHUH & SLATER 12a); body with fanlike (Figs 13c, h, i) or (1995) also separated these two taxa, on the clove-like (Figs 13a, b, f) setae intermixed basis of the less broad scutellum (and more with curly sericeous setae; pronotal, corial exposed abdominal connexiva) and reduced and connexival margins dentate (Fig. 11); hindwing intervannal vein in eurygastrines. male androconial glands present on abdom- We have not evaluated either of these char- inal sterna IV-VI (Figs 2d, e); ejaculatory acters, and uncritically support the separa- reservoir small, oval (Figs 14c, d); ventral tion of the taxa. It is noteworthy that many conducting canal straight; CAI membrane- odontotarsines and pachycorines have a re- ous, medially fused (Figs 14c, d); CAII duced metathoracic peritreme, or it is ab- strongly sclerotized, U-shaped (Figs 14c, d); sent entirely, with a distinct, laterally ori- CAIII present or absent (Figs 14c, d); vesi- ented ostiole, however these taxa are sepa- ca, elongate, arcuate or S-shaped (Figs 14c, rated on the absence of stridulatory vittae in d); female gonocoxae I tripartite; interlock- odontotarsines. ing rami absent; spermathecal fecundation The monophyly of the Odontotarsinae, canal short; spermathecal reservoir ellip- however, is contentious, and we know of no toid; and spermathecal bulb bilobed. character that defines them exclusively. The Description: Body oval; small species, above diagnosis is polythetic in format and males 4.51-5.37 mm, females 4.41-5.88 the group is retained largely as a ‘conven- mm; dorsum moderately convex; cryptozoic ience’ group. There are putative infra-subfa- colouring, yellow-brown to dark brown, milial which are worthy of examina- with patterned darker markings; dorsum tion. A group of genera related to Odon- with elongate clove-shaped (Figs 13a, b, f) toscelis, including those taxa with male se- to fanlike setae (Figs 13c, h, i), intermixed tose sternal glands (Table 3), and sometimes with curly sericeous setae (Figs 13a, e, h); have rounded lateral margins of the Prono- densely punctate. Head: semicircular, tum. This is largely a Palearctic group, but strongly deflexed (Figs 12a, b); lateral mar-

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Table 5: Morbora australis, M. hirtula, and M. schoutedeni: diagnostic measurements in margins convex, explanate, denticulate, an- millimetres. N = sample size. terolateral angles anteriorly produced, ex- Male Female tending beyond posterior margins of eyes N Mean±SD Range N Mean±SD Range (Figs 11a-c, 12a-c); posterior margins M. australis rounded to truncate (Figs 11a-c). Scutel- Length 1 5.37 - 4 5.40±0.22 5.27-5.73 lum: broad, U-shaped to subhemispherical Pronotal width 1 4.41 - 4 4.17±0.21 3.85-4.29 (Figs 11a-c), strongly convex in profile; Width between eyes 1 1.59 - 4 1.66±0.13 1.54-1.84 dentate corial and abdominal connexival Antennal segment length margins visible (Figs 11a-c). Thoracic ster- I 2 0.64 0.61-0.67 7 0.60±0.03 0.54-0.63 II 2 0.30 0.29-0.30 8 0.33±0.02 0.30-0.36 na: prosternal margins and anterior margin III 2 0.22 - 8 0.23±0.02 0.22-0.26 of proepisternum jointly raised, moderately IV 2 0.35 - 7 0.32±0.02 0.28-0.35 explanate (Fig. 12c). Thoracic pleura: ex- V 2 0.62 0.62-0.63 7 0.56±0.03 0.50-0.59 Rostral segment length ternal efferent system of metathoracic I 1 0.62 - 2 0.59 0.57-0.61 glands moderately developed (Fig. 12c, d); II 1 0.51 - 3 0.45 0.39-0.54 ostiole small; evaporative area of metatho- III 1 0.45 - 2 0.45 0.42-0.48 IV 1 0.34 - 2 0.33 - racic gland extending onto posterior por- Morbora hirtula tion of mesopleuron and beyond peritreme; Length 3 5.06 4.97-5.12 2 4.90 4.41-5.39 peritreme short, tongue-like, raised, and Pronotal width 3 3.64 3.58-3.72 2 3.55 3.31-3.80 wrinkled (Figs 12c, d). Pregenital abdomen: Width between eyes 3 1.53 1.47-1.57 2 1.58 1.54-1.62 broad, strongly convex; lateral margins Antennal segment length spinose; male abdominal SIV-VI with I 5 0.58±0.04 0.53-0.63 3 0.61 0.61-0.62 paired, suboval regions of androconial II(a) 5 0.33±0.03 0.30-0.36 2 0.36 0.35-0.36 II(b) 4 0.23±0.02 0.21-0.25 2 0.22 0.22-0.23 glands (Figs 2d-f). Male Genitalia: ventral III 3 0.40 0.35-0.42 2 0.42 0.41-0.42 surface of pygophore terminal in orienta- IV 5 0.65±0.03 0.62-0.70 2 0.64 0.63-0.65 tion (Fig. 12e), ventral margin entire (Fig. Labial segment length 14a); genital opening small to moderately I 1 0.82 - 1 0.85 - II 1 0.97 - 1 0.91 - sized (Figs 12f, 14a); parameres short, III 1 0.36 - 1 0.42 - crown small, hook-like, tip blunt (Fig. 14b); IV 2 0.37 0.36-0.38 1 0.45 - aedeagus broad; phallotheca conical, with- Morbora schoutedeni out processes, weakly sclerotized (Figs 14c, Length 2 4.74 4.51-4.97 5 5.47±0.34 5.00-5.88 d); ductus seminis proximalis thin, mem- Pronotal width 2 3.79 3.65-3.92 5 4.41±0.28 4.17-4.85 braneous (Figs 14c, d); ejaculatory appara- Width between eyes 2 1.40 1.35-1.45 5 1.70±0.09 1.59-1.84 tus simple (Figs 14c, d), with single, Antennal segment length I 4 0.54±0.02 0.51-0.57 9 0.68±0.06 0.61-0.76 straight, ventral conducting canal, without II(a) 4 0.25±0.03 0.22-0.28 9 0.28±0.03 0.23-0.33 convolutions; ejaculatory reservoir short, II(b) 4 0.23±0.01 0.23-0.24 10 0.22±0.01 0.21-0.24 suboval; dorsal conducting canal simple, III 4 0.29±0.02 0.27-0.31 10 0.32±0.04 0.24-0.39 not expanded; vesica broad, elongate, IV 4 0.49±0.03 0.46-0.52 9 0.55±0.04 0.46-0.61 weakly S-shaped (Figs 14c, d); CAI large, Labial segment length I 2 0.77 0.75-0.79 3 0.75 0.73-0.79 membraneous, medially fused, distally di- II 2 0.77 0.76-0.79 2 0.79 0.73-0.85 vided, without sclerotization (Figs 14c, d); III 2 0.36 - 2 0.41 0.36-0.46 IV 2 0.38 0.36-0.39 1 0.43 - CAII large, strongly sclerotized, U-shaped, with subdistal spur (Figs 14c, d); CAIII gins of jugae explanate, strongly denticu- membraneous, medially fused, distally free, late, surpassing apex of clypeus (Figs 12a, without sclerotization (Figs 14c, d). Female b); bucculae moderately tumescent, medial Terminalia: female paratergites IX S- margins subparallel (Fig. 12c); eyes small shaped; female gonocoxae I divided, tripar- (Figs 12a-c); antennae: AI elongate, either tite; interlocking rami absent. Spermathe- longest subequal or subequal to AIV; labi- ca: fecundation canal short; spermathecal um: reaching metasternum; either LI or LII reservoir weakly dilated, simple; and sper- longest segment, LIII & LIV small, either mathecal bulb bilobed, with proximal and distal flanges. subequal or LIII longest. Pronotum: sub- trapezoidal, moderately convex in profile Diversity and distribution: Morbora is (Figs 11a-c); anterior margin excavate; with endemic to continental Australia, com- broad, explanate paranotal lobes, lateral prised presently of three species.

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Included species: M. australis DISTANT 1899 Australia M. hirtula BERGROTH 1904 Australia M. schoutedeni BERGROTH 1904 Australia

Remarks: DISTANT (1899) first described Morbora, with SCHOUTEDEN (1904) and GROSS (1975) providing subsequent re- descriptions. MCDONALD &CASSIS (1984) redescribed the species, without providing any additional observations for the genus or comments on its relationships.

GROSS (1975) placed Morbora in the Odontotarsinae, on the basis of observations by Fred McDonald of the male genitalia, the latter worker suggesting that they resembled members of the Odontotarsinae. Our obser- vations confirm these findings, and have found further evidence that Morbora is nest- ed within the odontotarsines. As stated above, Morbora has male abdominal andro- conial glands, in common with some Pale- arctic odontotarsinae taxa (Table 3). More- over, we have found that some species of Psacasta (e.g., P. exanthematica SCOPOLI) have the first gonocoxae divided (bipartite), which is a putative homology to the tripar- tite GXI condition found in all the species of Morbora. Fig. 11: Habitus of Morbora species (a) M. australis (b) M. hirtula (c) M. schoutedeni. Scale bars = 1 mm. Morbora australis DISTANT 1899 (Figs 11a, 13a-c, 15, Table 5)

Morbora australis DISTANT 1899: 47 (n.sp.); SCHOUTEDEN 1904a: 87, pl. 5 fig. 10 (list; habi- Vestiture. Head and pronotum with tus); GROSS 1975: 90, fig. 21 (description; habi- dense distribution of erect, elongate, clove- tus); MCDONALD &CASSIS 1984: 548 (descrip- shaped setigerous punctures, intermixed tion; genitalia); CASSIS &GROSS 2002: 586 (cat- with curly sericeous setae (Figs 13a-c); alogue; localities); MOIR et al. 2003: 353 (West- scutellum with dense distribution of short, ern Australian record) fan-shaped setae, relatively narrow distally. Diagnosis: Morbora australis is recog- Structure. Antennae: AI longest seg- nised by the following combination of char- ment, little longer than AIV. Labium: LI acters: scutellum with clove-shaped and longest segment, LII & LIII roughly sube- short fan-shaped setae (Figs 13a-c); AI qual, LIV shortest segment. Pronotum: lat- longest segment; exocorium not expanded eral margins moderately expanded; antero- (Fig. 11a); CAI branched, lobe-like; CAII lateral angles not surpassing anterior margin elongate, sickle-shaped; and, CAIII absent. of eyes. Hemelytra: costal margins not great- Description: Body small, males 5.37 ly expanded laterally. Male Genitalia: CAI mm, females 5.27-5.73. branched, membraneous, lobe-like, without sclerotization; CAII elongate, sickle-shaped, Colouration. Dorsum yellowish-brown heavily sclerotized; CAIII absent; vesica to dark brown, with contrasting markings, heavily sclerotized, distally tapered. broad W-shaped yellow marking on posteri- or 1/2 of scutellum (Fig. 11a). Measurements. Table 5.

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Fig. 12: Scanning electron micrographs of key characters of Morbora australis (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ag = androconial glands; Ea = evaporative areas; Pe = peritreme; P(r) = right paramere. Scale bars = 1 mm.

Type material examined: Holotype, 1=, Peak Oliver, February 2001, DLWC WALCOL00149 Downs, B.M. Hem. Type No. 613, ‘Morbora aus- (AM); 2YY, Yelta, 30°27’14“S 148°41’42“E, I tralis DIST type’, ‘Dist. Coll. 1911-383’ (BMNH). Oliver, February 2001, DLWC WALCOL00025 Material examined: Queensland: 3YY, Walkers and WALCOL00065, ex pitfall trap (AM); 1Y, Creek campsite, 17°28’S 141°10’E, 28 July 1995, J Wentworth, May 1965, MJ Coulson (ANIC); Y Thompson, FN 997 (AM); 1 , Carnarvon Australian Capital Territory: 1Y, Black Moun- Range, March 1944, N Geary (AM); 1= 1Y, tain, 35°16’S 149°06’E, 600 m, March 1987, TA Clermont, viii-1929 KK Spence (AM); 1Y, Bluff Weir, JF Lawrence & W Dressler, ex flight inter- Range, near Biggenden, 21-iii-1975, H Frauca cept trap (ANIC); Southern Australia: 1Y, Ma- (ANIC); 1Y, Luster Creek, 8 km W by N Mt ree-Lyndhurst Road, J Upton, 15-ix-1972 (AN- Molloy, 21-22 May 1980, ID Naumann (ANIC); IC); Western Australia: 1Y, Kimberley District, New South Wales: 1= 1Y, Bundjalong National Park, 32°24’S 152°32’E, G Cassis, 15 November Emma Gorge Resort, 4.5 km N Gibb River Road, 1993, ex beach wash (AM); 2YY, 30°04’41“S 15°54’40“S 128°07’29“E, 4-13 June 1999, MR 148°56’53“E, I Oliver, February 2001, DLWC Gray, G Milledge & H Smith, pitfall trap, ex sa- WALCOL00286 and WALCOL00498 (AM); vanah woodland FN 14481 (AM). 1Y, Road Reserve, 30°11’42“S 148°54’54“E, I

324 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at ) broad fan-like i ) sericeous setae on pronotum ) clove-shaped setae on lateral e b ) fan-like and sericeous setae on head pronotum ( h ) sericeous and clove-shaped setae on pronotum ( a ( ) head and pronotum with sericeous clove-shaped setae ( d ( Morbora australis ) a-c Morbora hirtula ) ) dorsum with sericeous and fan-like setae ( species ( g ( d-f Morbora Morbora schoutedeni ) g-i ) short fan-like setae on scutellum ( c Abbreviations: cs = clove-shaped setae; fs fan-shaped setae. . : Scanning electron micrographs of vestiture ) clove-shaped setae on scutellum ( f setae on scutellum ( projections of pronotum ( Fig. 13

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Fig. 14: Male genitalia of Morbora schoutedeni (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAI = first conjunctival appendage; CAII = second conjunctival appendage; CAIII = third conjunctival appendage; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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Distribution: Morbora australis is widely habitus); KIRKALDY 1909: 265 (catalogue); GROSS distributed in Australia, mostly from arid or 1975: 91 fig. 22 (description; habitus); MCDON- semi-arid habitats. It is presently recorded ALD & CASSIS 2002: 548 (description; genitalia); from Queensland, New South Wales, Aus- CASSIS &GROSS 2002: 587 (catalogue) tralian Capital Territory, South Australia, Diagnosis: Morbora hirtula is recognised Victoria, Western Australia and the North- by the following combination of characters: ern Territory (Fig. 15). MOIR et al. (2003) dorsum with clove-shaped setae, intermixed reported this species from Western Aus- with curly sericeous setae (Figs 13d-f); AIV tralia. The Kimberley district record report- longest segment; exocorium not expanded ed in this work confirms the western distri- (Fig. 11b); CAI membraneous, lobe-like; bution of M. australis. CAII elongate, sickle-shaped, distally bifur- cate; and, CAIII lyre-shaped and heavily Host plant records and habitats: Morbo- sclerotized. ra australis has been collected primarily in pitfall traps or flight intercept traps, suggest- Description: Body small, males 4.97- ing it is epigaeic in its habits. The specimens 5.12, females 4.41-5.39. collected at Bundjalong National Park Colouration. Dorsum dark brown, with (NSW) were collected on beach wash, after contrasting markings, most notably with a violent thunderstorm at sea. Nothing is broad W-shaped marking on posterior 1/2 of known of its food-preferences. scutellum (Fig. 11b).

Remarks: DISTANT (1899) first described Vestiture. Dorsum with dense distribu- this species from Peak Downs in southeast tion of erect, elongate, clove-shaped setae, Queensland. GROSS (1975) redescribed it intermixed with curly sericeous setae, most- from a range of collections across eastern ly on head and pronotum; without fan- Australia, and South Australian and the shaped setae (Figs 13d-f). Northern Territory. He distinguished it from Structure. Antennae: AIV longest seg- M. hirtula, on the basis of its paler coloura- ment, little longer than AI. Labium: LII tion (and contrasting dorsal colour pattern- longest segment, LIII & LIV subequal in ing) and different vestiture (cf. Figs 13a-i). length. Pronotum: lateral margins moderate- MCDONALD &CASSIS (1984) redescribed ly expanded (Fig. 11b); anterolateral angles this species, and provided the first descrip- not surpassing anterior margin of eyes (Fig. tion (and illustration) of the male and fe- 11b). Hemelytra: costal margins not greatly male genitalia. They found that the aedea- expanded laterally. Male Genitalia: CAI gus has only two conjunctival appendages membraneous, lobe-like, without sclerotiza- (CAIII absent), which conclusively differ- tion; CAII elongate, sickle-shaped, heavily entiates it from M. hirtula. CASSIS & GROSS sclerotized, distally bifurcate, antler-like; (2002), in cataloguing this species, provided CAIII short, lyre-shaped, heavily sclerotized; additional distributional information. Our vesica heavily sclerotized, distally tapered. observations do not support the colour dis- Measurements. Table 5. tinction as mentioned above; we found that M. australis varies from yellow-brown to Material examined: Queensland: 3YY, 3 km dark brown, even within the same popula- NE of Mt Webb, 15°03’S 145°09’E, A Calder & J Feehan, 30 April-3 May, 1981, ANIC Berlesate tion. It is best determined by the distribu- 723, ex rainforest litter (AM, ANIC); New tion of short fan-like setae (Fig. 13c) partic- South Wales: 4 km NE Mt Wog Wog, 17 km SE ularly on the lateral margins of the prono- Bombala, 37°04’S 149°28’E, CR Margules, Octo- tum and across the scutellum. Some speci- ber 1991, ex pitfall (ANIC). mens also possess clove-shaped setae, as in Distribution: Morbora hirtula is widely M. hirtula, but the latter species always lacks distributed in eastern Australia, and reaches the fan-shaped setae. the Northern Territory to the west. It is known from locations in Queensland, New Morbora hirtula BERGROTH 1904 South Wales, and South Australia (Figs 11b, 13d-f, 15, Table 5) (Fig. 15). It has been recorded from Victoria Morbora hirtula BERGROTH 1904: 356 (n.sp.); and the Northern Territory, but without pre- SCHOUTEDEN 1904: 87, pl. 5 fig. 10 (description; cise locations.

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ocorium expanded (Fig. 11c); pygophore (Fig. 14a); parameres with short apical hook (Fig. 14b); CAI membraneous, lobe-like, bi- fid (Figs 14c, d); CAII elongate, sickle- shaped, distally bifurcate (Figs 14c, d); and, CAIII lobe-like, fused basally, membraneous (Figs 14c, d). Description: Small species, males 4.51- 4.97 mm, females 5.00-5.88 mm. Colouration. Dorsum dark brown, with contrasting markings, most notably with broad W-shaped marking on posterior 1/2 of scutellum (Fig. 11c). Vestiture. Dorsum with dense distribu- tion of erect, elongate setae, intermixed with curly sericeous setae, mostly on head and pronotum (Figs 13g-i); pronotum and particularly scutellum with dense distribu- tion of broad fan-shaped setae (Figs 13g-i). Structure. Antennae: AI longest seg- ment, little longer than AIV. Labium: LI & Fig. 15: Distribution of Morbora species. Host plants and habitats: Morbora hirtu- LII, and LIII & LIV subequal in length. la is also likely to be an epigaeic species; col- Pronotum: lateral margins greatly expanded lected in pitfall traps or through litter ex- (Fig. 11c); anterolateral angles surpassing traction. It is broadly distributed across anterior margin of eyes (Fig. 11c). Hemely- ecosystems, from open woodland to rainfor- tra: costal margins greatly expanded laterally est habitats. (Fig. 11c). Male Genitalia: pygophore with Remarks: Morbora hirtula is best identi- narrow genital opening (Fig. 14a); parameres fied by the uniform distribution of elongate, with a short apical hook (Fig. 14b); CAI large, membraneous, lobe-like, bifurcate, clove-shaped setae on the dorsum, and lacks without sclerotization (Figs 14c, d); CAII the fan-like setae that occur in its two con- elongate, sickle-shaped, heavily sclerotized, geners (cf. Figs 13a-i). Externally it is very with short distal bifurcation, antler-like (Figs similar to M. australis, but the male genitalia 14c, d); CAIII broad, membraneous, medial- differs significantly in having three con- ly fused; vesica broad, heavily sclerotized, junctival appendages. This is also the case weakly S-shaped (Figs 14c, d). in M. schoutedeni, but in M. australis the bi- furcation of CAII is more pronounced. Measurements. Table 5. Material examined: New South Wales: 1Y, Morbora schoutedeni BERGROTH 1904 Baraba, 30.115S 148.790 E, I Oliver, February (Figs 11c, 13g-i, 14, 15, Table 5) 2001, Site WALCOL00162, ex pitfall trap, (AM); 1Y, Nerrub, 30°04’41“S 148°56’53“E, I Morbora schoutedeni BERGROTH 1904: 356 (n.sp.); Oliver, February 2001, Site WALCOL00285, ex SCHOUTEDEN 1904: 87 (list, habitus); KIRKALDY pitfall trap (AM); 1Y, Yetta, 30°27’04“S 1909: 265 (catalogue); GROSS 1975: 92 (descrip- 148°41’35“E, I Oliver, February 2001, Site WAL- tion, habitus); MCDONALD & CASSIS 1984: 548 COL00068, ex pitfall trap (AM); 1Y, Womba, (description); CASSIS & GROSS 2002: 586 (cata- 30°23’60“S 148°41’53“E, I Oliver, February 2001, logue) Site WALCOL00568, ex pitfall trap (AM); 2==, Diagnosis: Morbora schoutedeni is recog- Bogan River, J Armstrong (AM; Queensland: = nised by the following combination of char- 1 , Clermont, November 1929, KK Spence (AM); Western Australia: 2YY, Pilbara Dis- acters: dorsum with broad fan-shaped setae, trict, Hammersley Station, Kaengaenarina well, intermixed with curly sericeous setae (Figs 500 m from well, 22°15’44“S 159°48’38“E, 14-22 13g-i); AI longest antennal segment; prono- April 2005, S Lassau, M Elliott, L Kampen & M tal lateral margins expanded (Fig. 11c); ex- Bulbert, PILB126/05P (AM).

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Distribution: This species is broadly dis- Agonosoma LAPORTE 1832 tributed in Australia (New South Wales, (Figs 16, 17, 18, Table 6)

Northern Territory, Queensland, South Eurygaster (Agonosoma) LAPORTE 1832: 62 Australia) in semi-arid and arid areas (Fig. Agonosoma: SCHOUTEDEN 1904: 67 (generic sta- 15). In this work we record it from Western tus; description); AMYOT & SERVILLE 1843: 44 Australia (Pilbara District). It is sympatric (description); STÅL 1867: 494 (description); with M. australis at a number of sites in east- KIRKALDY 1909: 278 (catalogue); EGER 1987: 339 ern Australia (NSW: Yelta, Nerrub; QLD: (diagnosis); PALEARI 1992: 505 (description; revi- Clermont). sion) Agonocoris BERGROTH 1891: 235 (unnecessary Host plants and habitats. Morbora nom. nov.) schoutedeni is a ground-dwelling species, Type species: Agonosoma flavolineata LAPORTE which has been mostly collected in pitfall 1832, monotypy traps. As with the other Morbora species, Diagnosis: Agonosoma is recognised by nothing is known of its food preferences. the following combination of characters:

Remarks: BERGROTH (1904) first de- body elongate-ovoid (Figs 16a, b), moder- scribed this species from Townsville ately convex dorsum; body mostly glabrous (Queensland). GROSS (1975) redescribed it, (Figs 16a, b); densely punctate with small and diagnosed it in part by the reportedly punctures (Figs 16a, b, 17a); head large, ju- subequal AII(a) (as third segment) and gal margins weakly sinuate (Fig. 17a), thick- AII(c) (as fourth segment). In our work, we ened and rounded in profile; bucculae later- have measured specimens of all three al margin anteriorly compressed (Fig. 17b); species, and found this character to not eyes small, contiguous with pronotum, over- have diagnostic value, with continuous vari- lapping anterolateral angles of pronotum ation existing, such that AII(b) can be (Fig. 17a); anterior margin of proepisternum longer than AII(a), and overlaps in the moderately explanate (Fig. 17b); pronotum ranges of these characters were found with trapezoidal, anterior margin weakly exca- the other Morbora species. This species is vate, anterolateral margins elongate, strong- distinctive having the broad fan-like setae ly divergent, posterior margin truncate (Figs that are very densely distributed on the 16a, b); scutellum strongly declivent poste- pronotal disc and scutellum (Figs 13g-i). riorly, rounded distally; mesepimeron with The conjunctival appendages are also dis- evaporative areas (Fig. 17c); external effer- tinctive, with medially fused CAIII, and ent system well-developed, peritreme sickle- CAII with a short distal bifurcation. shaped, medially sulcate, anteriorly re- curved, evaporative area extending beyond Subfamily Pachycorinae peritreme (Figs 17c, d); abdominal SV-VI Remarks: It is beyond the scope of this with stridulatory vittae (Figs 17e, f); and, work to review the Pachycorinae, which are SVII enlarged in both males and females not native to Australia; we only reviewed covering terminalia. (Fig. 17d). the introduced species, Agonosoma trilinea- Diversity and distribution: Agonosoma tum in any detail. This subfamily is largely has four species, which are endemic to the found in the Western Hemisphere, with on- tropics of the Neotropical zoogeographic re- ly Deroplax MAYR and Hotea occurring in gion (KIRKALDY 1909; PALEARI 1992). the Old World (KIRKALDY 1909; SCHUH & Included species: SLATER 1995). KUMAR (1964) placed the Australian species, Tectocoris diophthalmus, A. dohrni SCHOUTEDEN 1903 Mexico in the Pachycorinae, on the basis of the A. bicolor WESTWOOD 183 Brazil ejaculatory apparatus; an action not fol- A. flavolineatum LAPORTE 1832 Brazil, lowed by subsequent authors. Pachycorines Guyana, Venezuela are primarily diagnosed by the presence of A. trilineatum (FABRICIUS 1781) stridulatory vittae and tibial plectra; charac- Colombia, Brazil, Guyana, ters that are thought to be exclusive to the Panama, Suriname, Venezuela, West Indies, Australia (introduced) Pachycorinae (e.g., SCHUH & SLATER 1995).

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Agonosoma trivittata fabricii KIRKALDY 1909: 278 (unnecessary nom. nov.) Diagnosis: Agonosoma trilineatum is recognised by the following combination of characters: males chocolate brown, with three longitudinal yellow stripes (Fig. 16a); females dimorphic, as in male, or mostly yel- low-brown, with pair of submedial spots on pronotum and scutellum (Fig. 16b). Colouration. Males monomorphic, dor- sum dark chocolate brown with three longi- tudinal yellow stripes (Fig. 16a); females di- morphic, either as in male, or pale brown with black spots (Fig. 16b). Head: jugal mar- gins black, subjugal regions of frons orange, large fuscous triangular marking from poste- rior margin of head to near anteclypeus, with medial yellow stripe; underside of head mostly iridescent green, with posterior re- Fig. 16: Habitus of introduced Remarks: Agonosoma was described as a gion orange; AI orange, sometimes tip em- Pachycorinae species (a-b) Agonosoma subgenus of Eurygaster by LAPORTE (1832). browned, remainder of segment fuscous; LI- trilineatum (a) = (b) Y. Scale bar = 1 mm. SCHOUTEDEN (1904) redescribed the genus II mostly dusty stramineous, LIII-LIV fus- and provided a comprehensive synonymy cous; Pronotum: spotted morph with 6 for the species. EGER (1987) in revising Tiri- spots, pairs on anterolateral angles, subme- dates STÅL, recognised a group of genera dially on anterior margin, and submedially (Agonosoma, Crathis STÅL, MAYR, below callosite region. Thoracic pleura: an- Lobothyreus MAYR, DALLAS and terior edge of proepisternum yellow, remain- Tiridates) with enlarged abdominal SVII (in der of pleura fuscous, with green iridescent both sexes), concealing the genitalia. Fur- tinge, intermixed with yellow posterior mar- ther, EGER (1987) provided differential diag- gin of mesepimeron and peritreme and part noses for Agonosoma and Tiridates, placing of supracoxal lobes. Legs: coxae yellow; most importance on the external efferent femora orange; tibiae fuscous in stripped system of the metathoracic glands, and sec- morph, or mostly orange with anterior sur- ondary diagnostic value to the body and face fuscous; tarsi fuscous. Abdominal Ven- head shape. PALEARI (1992) redescribed ter: ground colour yellow-orange, with later- Agonosoma and the constituent species, but al margins dark brown and green irides- did not detail the genitalia. cence, SV-SVI stridulatory vittae dark brown, SII mostly dark brown, anterior mar- Agonosoma trilineatum (FABRICIUS gins of SIII-VII dark brown. 1781) (Figs 16, 17, 18, Table 6)

Cimex trilineatus FABRICIUS 1781: 341 (n. sp.) Texture. Dorsum, underside of head and Cimex sexpunctatus FABRICIUS 1781: 339 (n.sp.); thoracic pleura punctate (Figs 16a, b, 17b). PALEARI 1992: 513 (synonymy) Vestiture. Body glabrous. Antennae: Cimex trivittatus PANZER 1798: 111 (n.sp.); AI-AII(a) almost glabrous, AII(b)-AIV PALEARI 1992: 513 (synonymy) with dense distribution of short, simple, se- virgatus GERMAR 1839: 102 (n.sp.); PALEARI 1992: 513 (synonymy) mi-erect setae; femora with sparse distribu- Agonosoma quadiguttatum SIGNORET 1851: 330 tion of short setae; ventral surface of tibiae (n.sp.); WALKER 1867: 59 (list); PALEARI 1992: with dense distribution of short, semi-erect 513 (synonymy) setae. Agonosoma trivittatum: SIGNORET 1851: 330 (new Structure. Antennae: AII(a) smallest combination); DALLAS 1851: 42 (list) Agonosoma trilineatum: STÅL 1870: 13 (new com- segment, AIV longest segment, > AIII, bination); SCHOUTEDEN 1904: 68 (synonymy); AII(b) little longer than AII(a), AII(b) PALEARI 1992: 505 (redescription) subequal to AI; AIII and AIV flattened.

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Fig. 17: Scanning electron micrographs of key characters of Agonosoma trilineatum (a) Head, dorsal view (b) Head, lateral view (c) External efferent system of metathoracic glands, ventral view (d) male pygophore, dorsal view (e) Stridulatory areas, pregenital abdominal SV- SVI (f) Substructure of stridulatory area. Abbreviations: Ea = evaporative area; Pe = peritreme; P(r) = right paramere; S = stridulatory area; SVII = seventh abdominal sternite, dorsal view. Scale bars = 0.1 mm.

Labium: reaching abdominal SIII, LII (Fig. 18c). Female Terminalia: paratergites longest segment. Male Genitalia: pygophore VIII small, subtriangular, tapered medially; with narrow genital opening, laterally setose paratergites IX large, subelliptoid, medial (Fig. 17d, 18a); parameres with short, broad margins rounded; gonocoxae I bipartite, stem and angulate, elongate crown (Fig. posterior, digitiform branch, anteriorly sub- 18b); aedeagus with sclerotized, broad duc- triangular, moderately-sized. Spermatheca: tus seminis proximalis, bounded by ligamen- fecundation canal short; reservoir very tory tubule (Fig. 18c); ejaculatory apparatus weakly dilated, thickened; pump elongate, with ventral conducting canal broad, not with proximal and distal flanges present; convoluted (Fig. 18c); ejaculatory reservoir bulb small, round. elongate, subreniform (Fig. 18c); ductus Measurements. Table 6. seminis distalis broad, scerlotized (Fig. 18c, Material examined: Venezuela: 5== 11YY, d); vesica membraneous, denticulate (Fig. Falcon State, Paraguana Peninsula, 15 December 18c, d); CAI and CAIII absent (Fig. 18c); 1999, R Segura & TA Heard, ex Jatropha gossypi- CAII broad, antler-like, distally denticulate folia, quarantine colony (AM).

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Table 6: Agonosoma trilineatum: diagnostic measurements in millimetres. N = sample size. body (e.g., Figs 19a-d, 28a); ovoid (e.g., Figs 24b, 37) to elongate-ovoid (Figs 28a, b), Male Female sometimes strongly tapered towards termi- N Mean±SD Range N Mean±SD Range nalia; body weakly (e.g., Fig. 28a) to heavi- Length 9 8.70±1.05 7.36-10.42 9 9.72±0.67 9.14-11.07 Pronotal width 9 5.40±0.43 4.84-5.93 9 6.01±0.34 5.53-6.62 ly punctate (Figs 24a, c-f); body brilliant, Width between eyes 9 2.42±0.14 2.22-2.62 9 2.59±0.14 2.27-2.77 aposematically coloured, often with irides- Antennal segment length cent blue, green, orange colouration, often I 18 0.75±0.05 0.70-0.85 17 0.82±0.07 0.75-1.00 with contrasting markings (e.g., Figs 19a-d, II(a) 18 0.59±0.06 0.50-0.68 16 0.62±0.08 0.48-0.75 24a, c-f, 28a-e); head subtriangular to subo- II(b) 18 0.77±0.06 0.65-0.85 16 0.86±0.07 0.78-0.95 III 18 1.37±0.07 1.25-1.48 12 1.50±0.07 1.35-1.58 val (e.g., Fig. 31a); thoracic sterna flat (e.g., IV 14 1.84±0.11 1.65-2.00 10 1.85±0.05 1.78-1.90 Figs 20c, 29c, 31c); anterior margin of Labial segment length proepisternum weakly explanate (e.g., Figs I 9 2.15±0.05 2.05-2.20 9 2.22±0.11 2.05-2.40 20c, 31c); efferent system of metathoracic II 9 2.22±0.06 2.15-2.33 9 2.42±0.11 2.25-2.55 III 8 0.85±0.11 0.73-1.00 8 0.90±0.07 0.85-1.03 glands well-developed (e.g., Figs 20d, 25d, IV 8 0.97±0.05 0.90-1.05 8 1.09±0.04 1.03-1.15 31d); peritreme mostly subreniform (e.g., Figs 20d, 39d, 42d), sometimes large, subtri- Distribution: This species was intro- angular (e.g., Figs 25d, 29d), or obovate duced to southeast Queensland, and natu- (e.g., Figs 31d, 38c); males without abdomi- ralised populations are as yet to be recovered nal androconial or setose sternal glands; pre- (Heard pers. comm.). It is naturally known genital abdomen in both sexes without from Panama, Colombia, Venezuela, Suri- stridulatory vittae; ventral surface of py- nam, Grenada and Trinidad (PALEARI gophore caudally (e.g., Figs 32a-h) or ven- 1992). trally oriented (e.g., Figs 29e, 30e); para- Host plants and biology: Agonosoma tri- meres hook-shaped (e.g., Figs 22b, 40b); lineatum was introduced as a biocontrol ejaculatory apparatus usually well-devel- agent of Jatropha gossypifolia (see above). oped, with convoluted ventral conducting canal and oval to subelliptoid ejaculatory Remarks: PALEARI (1992) redescribed reservoir (e.g., Figs 30c, d, 36c, d, 40c, d); this species, highlighting the colour dimor- CAI usually absent, rarely present (e.g., Figs phism, external morphology and distribu- 22c, d); CAII usually bifurcate, with lobal tion. It is distinguished from all the native sclerites (e.g., Figs 22c, d, 36c, d, 40c, d), jewel bugs of Australia, on colour and CAIII undivided, often U-shaped (e.g., Figs colour patterning alone. It is easily diag- 26d, 30d, 43d) or divided (e.g., Figs 22c, d); nosed by the enlarged abdominal SVII of spermathecal fecundation canal short or both sexes (covering the terminalia) (Fig. long; and, spermathecal reservoir oval. 17d, male), the membraneous vesica (Figs 18c, d), and thickened and narrow sper- Remarks: There is no unique character mathecal reservoir. state defining the Scutellerinae, and most of the above-mentioned features are also true Subfamily Scutellerinae for the family. They are the most brilliantly

Pentatomidae, Scutellerina, Scutelleraria STÅL coloured of all the scutellerids, but numerous 1872: 32 (new suprageneric taxon) Pachycorinae also exhibit colouration con- Pentatomidae, Scutellerinae, Scutelleraria: sistent with models of warning colouration. SCHOUTEDEN 1904: 13 (description) The male and female genitalia are also simi- Pentatomidae, Scutellerinae, Scutellerini: lar to taxa in other scutellerid subfamilies, KIRKALDY 1909: 289 (catalogue) such as the Elvisurinae and Sphaerocorinae, Pentatomidae, Scutellerinae, Scutellerini: LE- although the development of the ventral STON 1952d: 13 (new suprageneric classification) conducting canal of the ejaculatory appara- Scutelleridae, Scutellerinae: MCDONALD 1966: tus is putatively greatest in the scutellerines, 67, 68 (new suprageneric classification) such as in Cantao parentum (Figs 30c, d). Scutelleridae, Scutellerinae: MCDONALD & CAS- Many species have a U-shaped, and basally SIS 1984: 550 (Australia) fused CAIII (e.g., Figs 26d, 30d, 43d), but Diagnosis: The Scutellerinae are recog- there are significant variations, particularly nised by the following combination charac- in Calliphara (Fig. 22d). Scutellerines can be ters: small (e.g., Figs 24b, 37) to very large separated from the Pachycorinae, and Odon-

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Fig. 18: Male genitalia of Agonosoma trilineatum (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) apex of aedeagus, ventral view. Abbreviations: C = crown of right paramere; DS(D) = ductus seminis distalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; F = flange at base of crown of paramere; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; V = vesica; VM = ventral margin of genital opening.

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abdominal sterna (recurved anteriorly) and the presence of two intervannal veins in the hindwing. Neither of these characters is considered exclusive to this conception of the taxon. We have retained the scuteller- ines in the narrow sense, despite it being a ‘convenience’ group, as it helps in distin- guishing the colourful Australian scutellerid species, pending a more in-depth review of the issue.

Calliphara GERMAR 1839 (Figs 19, 20, 21a-c, 22, 23)

Calliphara GERMAR 1939: 122 (gen. nov.); HER- RICH-SCHAEFFER 1839: 19, 20, 22, 58 (descrip- tion); STÅL 1865: 33 (key); STÅL 1873: 16 (de- scription); LETHIERRY &SEVERIN 1893: 23 (cata- logue); SCHOUTEDEN 1904: 24, 31 (description); KIRKALDY 1909: 297 (catalogue); MCDONALD 1961: 183 (transferred to Sphaerocoraria); LYAL 1979 (revision); MCDONALD &CASSIS 1984: 561 (description); CASSIS & GROSS 2002: 587 (cata- logue) Lamprophara STÅL 1865: 34 (gen. nov.); LYAL 1979: 154 (synonymy) Calliphara (Chrysophara) STÅL 1873: 17 (subgen. nov. ); LYAL 1979: 154 (synonymy) Type species: Calliphara: Cimex imperialis FABRI- CIUS 1775, subsequent designation, KIRKALDY 1909: xxxv; Lamprophara: Calliphara (Scutellera) bifasciata WHITE 1839, monotypy; Calliphara (Chrysophara): Tetyra excellens BURMEISTER 1834, subsequent designation KIRKALDY 1909: xxxv. Diagnosis: Calliphara is recognised by the following combination of characters: glabrous; lateral margins of jugae deeply ex- cavate (Fig. 20a), rounded in profile (Fig. 20b); AII(a) shortest segment; forewing tip extending beyond scutellum (Figs 19a-d); proepisternum with short anterior keel (Fig. 20c); broad subreniform, raised peritreme; Fig. 19: Habitus of Australian Calliphara totarsinae + Tectocorinae, in lacking stridu- CAI divided (Figs 22c, d); CAII(L) promi- = species (a) Calliphara dimidiata cruenta, latory vittae and androconial/setose sternal (b) Calliphara imperialis, = (c) Calliphara nent digitiform lobal sclerite, CAII(M) nobilis, = (d) Calliphara regalis, =. glands respectively. They can be separated membraneous with bifid lobal sclerites (Figs Scale bar = 1 mm. from the Elvisurinae, Odontotarsinae and 22c, d); vesica often with apical process dis- Sphaerocorinae superficially by their apose- tal to secondary gonopore (Figs 22c, d); and, matic colouration (cf. dull colouration), al- spermatheca with short proximal fecunda- though even in the latter subfamily some tion canal; and, spermathecal reservoir taxa exhibit moderately striking colouration, large, oval. as in Sphaerocoris BURMEISTER. Description: Body elongate to elongate-

SCHUH & SLATER (1995) adopted LE- ovoid (Figs 19a-d); Large species, males 13.00-19.00 mm, females 14.00-20.00 mm. STON’s (1952d) definition of the Scutelleri- nae (including the Elvisurinae and Sphaero- Colouration. Body iridescent coloura- corinae), but on the basis of the emarginate tion, either orange-brown, blue, green or

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Fig. 20: Scanning electron micrographs of key characters of Calliphara imperialis (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ea = evaporative area; Go = genital opening; Pe = peritreme; P(r) = right paramere. Scale bars = 1 mm.

black, often with patterned, contrasting rior margin of metasternum, LII longest seg- markings (Figs 19a-d). ment, LIII and LIV roughly subequal. Pronotum: broadly trapezoidal (Figs 19a-d); Texture. Body impunctate to moderate- anterior margin weakly excavate; anterolat- ly punctate, with shallow punctures (Figs 19a-d). eral margins rectilinear, strongly divergent, carinate in profile (Figs 19a-d); posterolater- Vestiture. Body mostly glabrous. al margins weakly convex (Figs 19a-d); pos- Structure. Head: large, subtriangular, terior margin weakly excavate to rectilinear moderately declivent; strongly produced in (Figs 19a-d). Scutellum: broadly U-shaped, front of eyes (Fig. 20a); lateral margins of ju- strongly convex (Figs 19a-d), base of exoco- gae strongly excavate (Fig. 20a), rounded in rium, membrane tip and abdominal connex- profile (Fig. 20b); bucculae parallel-sided iva visible. Thoracic pleura: anterior margin (Fig. 20c). Antennae: not greatly elongate; of proepisternum moderately explanate (Fig. AII(a) often very short (Figs 20a, b); 20c); mesepimeron with evaporative areas; AII(b)-AIV flattened, roughly subequal in external efferent system of metathoracic length, with AIV longest, and AIII a little glands well-developed; peritreme broad, longer than AII(b). Labium: reaching poste- sickle-shaped, shallowly sulcate medially,

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apex anteriorly directed (Figs 20c, d); evap- lands and New Caledonia. Within Australia, orative areas extending beyond peritreme most species of Calliphara are found in north- (Fig. 20d). Pregenital Abdomen: mostly ern Queensland, with a few species extending posterolateral angles of connexiva acumi- as far south as northern New South Wales, nate. Male Genitalia: pygophore large, sub- and west to the Northern Territory. LIS & oval, dorsal in orientation (Figs 20e, f, 22a), SKÓRKA (1996) transferred C. bipunctata to most often with dorsal and ventral patches Notocalliphara LYAL. of setae (Figs 20f, 22a); parameres with Included species: crown hooked, apex blunt (Fig. 22b); aedea- C. bifasciata WHITE 1839 Fiji, Samoa gus: ductus seminis proximalis narrow, mem- C. billardierii (FABRICIUS 1803) Australia, braneous, tubelike (Figs 22c, d); ejaculatory Indonesia, New Guinea, apparatus with short, paired ventral con- Solomon Islands ducting canal, weakly convoluted (Figs 22c, C. caesar (VOLLENHOVEN 1863) Indonesia, d); ejaculatory reservoir small, suboval; dor- New Guinea sal conducting canal not expanded (Figs C. dimidiata dimidiata (DALLAS 1851 22c, d); vesica short, often with distal scle- New Guinea rotized process (Figs 22c, d); CAI divided C. dimidiata cruenta STÅL 1873 Australia post-thecal margin, either basally membra- C. dimidiata fasciata (WALKER 1867) neous, CAI(V) with short conical lobal New Guinea sclerite (Figs 22c, d) or uniformly sclero- C. excellens excellens (BURMEISTER 1834) tised, and CAI(D) greatly enlarged, subrec- Philippines, Indonesia tilinear to arcuate, sclerotised process (Figs C. excellens coelestis TAEUBER 1929 22c, d); CAIII bifid, with outer arcuate, Philippines sclerotised process, and medial membrane- C. excellens speciosa (WHITE 1842) ous lobe, often with bifid lobal sclerite (Figs Philippines 22c, d). Female Terminalia: paratergites IX C. imperialis (FABRICIUS 1775) Australia small, suboval; gonocoxae I subtriangular, C. lanceolata DISTANT 1903 outer surface flat, posterior margin weakly Indonesia (Timor) excavate; interlocking rami present. Sper- C. munda STÅL 1866 China matheca: proximal fecundation canal short; C. nobilis (LINNAEUS 1763) Australia, reservoir oval to barrel-shaped, with ribbed Burma, China, Indonesia, texture; distal fecundation canal with prox- Malaysia, Philippines, Taiwan imal and distal flanges; and, bulb oval. C. placida placida BREDDIN 1905 Diversity and distribution: Calliphara New Guinea comprises 15 species, five (dimidiata, excellens, C. placida scintillans BREDDIN 1905 placida, praslinia and regia) of which are divid- New Guinea ed into subspecies. The genus occurs primari- C. praslinia praslinia (GUÉRIN-MÉNEVILLE ly east of Wallace’s Line in southeast , 1838) Australia, Indonesia, with most species occurring in Indonesia, New Ireland, Solomon New Guinea and Australia. The genus occurs Islands, Vanuatu as far north as China (C. munda and C. no- C. praslinia admiraltyensis KIRKALDY 1909 bilis) and extends through the east Melane- Admiralty Islands sian Islands, and reaches most eastward to C. regalis (FABRICIUS 1775) Australia, Samoa, with inclusions in Vanuatu, New Indonesia, New Caledonia, Caledonia and Fiji. The genus is represented New Guinea, Solomon Islands in Australia by six species, with only C. im- C. regia regia (WESTWOOD 1837) perialis and the subspecies C. dimidiata cruen- Indonesia (Timor) ta, endemic. Calliphara nobilis is re-established C. regia allorensis LEHMANN 1920 as an Australian species, based on a collec- Indonesia (Timor) tion from the Northern Territory. The re- C. regia timorensis LEHMANN 1920 maining Australian species are widespread in Indonesia (Timor) the Australian zoogeographic region, known C. solomonensis LYAL 1979 Solomon Islands in both Indonesia and New Guinea, and in C. vollenhoveni LYAL 1979 some instances also in the east Melanesian Is- Bismarck Archipelago

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Species excluded from the Australian (2002) this species was recorded from the Fig. 21: Colouration of Abdominal Venter in Australian Scutellerinae (a) Calliphara fauna: CASSIS & GROSS (2002) included Northern Territory, Queensland and New dimidiata cruenta (b) Calliphara imperialis two species of Calliphara, C. billardierrii and South Wales. On re-examination of these (c) Calliphara nobilis (d) Calliscyta stalii (e) C. praslinia, in the Australian fauna. These collections, we have not found any speci- Cantao parentum (f) Lampromicra aerea are now considered to be erroneous records. mens that belong to this species. A speci- (g) Lampromicra regia (h) Lampromicra The Australian records of Calliphara men identified as C. praslinia from the senator (i) Scutiphora pedicellata. praslinia were based on misidentification of South Australian Museum has ‘N.T.’ as the museum collections. In CASSIS & GROSS label data; this however cannot confidently

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Fig. 22: Male genitalia of Calliphara regalis (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) apex of aedeagus, ventral view. Abbreviations: C = crown of paramere; CAI(V) = ventral branch of first conjunctival appendage; CAI(D) = dorsal branch of first conjunctival appendage; CAII(L) = lateral branch of second conjunctival appendage; CAII(M) = medial branch of second conjunctival appendage; CAIII = third conjunctival appendage; DS(D) = ductus seminis distalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; SP = setose patches of male genital opening; VCC = ventral conducting canal; Vp = distal process of vesica; VM = ventral margin of genital opening.

338 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at be regarded as a contraction for the ‘North- ern Territory’. In addition, this specimen is more likely to be C. excellens. In addition, there are various collections of Calliphara from the Melanesian zoogeographic subre- gion, which have been identified as C. praslinia. These specimens superficially re- semble C. praslinia in terms of the coloura- tion patterns. However, an examination of the male genitalia, indicate that they are very similar to C. regalis.

CASSIS & GROSS (2002) recorded Cal- liphara billardierii as an Australian species, based on KIRKALDY’s (1909) distributional descriptor of ‘Tropical Queensland’ for this species, but without reference to a precise locality. We have investigated all known collections of this species, and not found any specimens of C. billardierri, and there- fore exclude it from the Australian biota.

Remarks: LYAL (1979) reviewed Cal- liphara, describing new taxa, establishing new synonymies, and detailing the male genitalia. genera exist in the morphology of the ejacu- Fig. 23: Distribution of Australian Calliphara species. He subdivided the genus into four informal latory apparatus and conjunctival ap- species groups (C. excellens, C. praslinia, C. pendages. Calliphara has the following attrib- caesar and C. dimidiata) based on characters utes: simple male ejaculatory apparatus with of the male genitalia, particularly the devel- simple and short ventral conducting canal opment of the conjunctival appendages, and (Fig. 22d), vesica with distal process (Fig. length and shape of the vesica. 22d), CAI with enlarged digitiform lobal sclerites (Figs 22c, d), CAII distally with bi- LYAL (1979) proposed that Calliphara fid lobal sclerite, lateral branch of CAIII was most closely related to with long sclerotized, digitiform process (Figs HAHN and Lampromicra, but without explic- 22c, d), and medial branch of CAIII with bi- it morphological hypotheses. On external fid lobal sclerite (Fig. 22c). We have not ex- characters, Calliphara and Lampromicra amined any specimens of Chrysocoris, but closely resemble each other on the basis of published descriptions of the male aedeagus the following characters: sinuate lateral (TSAI et al. 2004), indicate a closer relation- margins of the jugae (cf. Figs 20a & 39a), ship of this genus with Lampromicra than explanate anterior margin of the proepister- Calliphara, in that the ejaculatory apparatus num (cf. Figs 20c & 39c), mesepimeron has an elongate and highly convoluted ven- with evaporative bodies (cf. Figs 20d & tral conducting canal. 39d), large metathoracic gland ostiole (cf. Figs 20d & 39d), subreniform peritreme (cf. Calliphara dimidiata cruenta Figs 20d & 39d), suboval male pygophore STÅL 1873 (Figs 19a, 21a, 23)

(cf. Figs 20f & 39f), dorsal margin of genital Calliphara cruenta STÅL 1873: 17 (n.sp.); LETHIER- opening of male pygophore with broad se- RY & SEVERIN 1893: 23 (catalogue); SCHOUTEDEN tose patch (cf. Figs 20f & 39f), small suboval 1904: 32 (list); KIRKALDY 1909: 297 (catalogue); female paratergites IX, and angulate posteri- MCDONALD 1963a: 28 (male genitalia) or margin of gonocoxae I. Calliphara dimidiata cruenta: LYAL 1979: 170 (as subspecies); MCDONALD & CASSIS 1984: 563, Calliphara differs externally from Lam- Fig. 75 (description; spermatheca) promicra by the larger body and very short Calliphara dimidiata cruentata: CASSIS & GROSS second antennal segment (Figs 20a, b). The 2002: 589 (incorrect subsequent spelling; cata- most significant differences between the two logue)

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Diagnosis: Calliphara dimidiata cruenta is distally moderately sclerotised, tapered dis- recognised by the following characters: tally; CAIII(D) broad sickle-shaped, with head, pronotal callosite region and posterior tapered apex, sclerotized; CAIII(V) absent; 1/2 of scutellum iridescent green; remainder vesica short. Female Terminalia: parater- of pronotum and scutellum orange (Fig. gites VIII large, subtriangular, medially sep- 19a); posterior angles of abdominal SIV-VII arated; paratergites IX smaller, subelliptoid; spinose (Fig. 21a); sternites of pregenital ab- gonocoxae I large, subtriangular, flap-like. domen orange (Fig. 21a); terminalia irides- Measurements. MCDONALD &CASSIS cent green (Fig. 21a); legs uniformly irides- 1984: Table 6. cent dark blue; labium reaching posterior margin of abdominal SIII; CAI claw-shaped; Material examined: Queensland: 1= 1Y, Leo CAII(M) with bulb-shaped apical process; C[ree]k track, approx. 300 m, McIlwraith Range, == CAIII bifid; and, vesica short. 11 January 1990, MS & BJ Moulds (AM); 2 2YY, Kelsal Collection (BMNH); 1Y, Cape Description: Body elongate-ovoid; large, York, Dämel, Australia Coll. 92-6 (BMNH); Tor- males 13-16 mm, females 14-17 mm. res Strait Islands: 3YY, 77-29 (BMNH); 2YY, Murray Is., 78-66 (BMNH). Colouration. Body bicoloured (Fig. Distribution: CASSIS & GROSS (2002) 19a), head, anterior and lateral margins of reported the known records of this species, pronotum, posterior 1/2 of scutellum, tho- from northern Queensland, the Torres Strait racic pleura and sterna, and abdominal ter- Island and Papua New Guinea (Fig. 23). minalia bluish-fuscous, often with purplish hue and green iridescence, posterior 1/2 of Host plant and biology: No host plant pronotum, anterior 1/2 of scutellum, and has been recorded for this species.

pregenital abdominal venter orange (Fig. Remarks: LYAL (1979) established Cal- 21a); antennae, labium and legs uniformly liphara dimidiata in its own species-group, and fuscous. also erected C. cruenta as a subspecies of C. Texture. Body mostly smooth, with dimidiata. He also delineated three subspecies scattered shallow punctures on dorsum. on external colour characteristics, with the abdominal sterna orange to red basally in the Vestiture. Dorsum glabrous; abdominal New Guinea subspecies (nominotypical sub- venter with sparse distribution of pale, sim- species and C. dimidiata fasciata) compared ple setae. to C. dimidiata cruenta, with darker sternites. Structure. Eyes: small, about 1/3 length In the Australian subspecies, the CAIII(V) of head; Antennae: AII(a) little longer than is absent and the CAII(M) has only one sub- AI. Labium: reaching abdominal sterna IV. distal spine. The subspecific status of these Pregenital Abdomen: posterior angles of ab- taxa was not evaluated in this work, al- dominal SVI-VII with acuminate spines though it is noteworthy that the bulb-like (Fig. 21a). Male Genitalia: ventral margin process of CAIII(M) is unique to all the sub- of male pygophore deeply emarginate; geni- species of C. dimidiata. MCDONALD & CAS- tal opening of pygophore with dorsal and SIS (1984) redescribed the species, including ventral patches of setae; ejaculatory appara- the first description of the female genitalia. tus small; ventral conducting canal, with two pairs of convolutions; ductus seminis Calliphara imperialis (FABRICIUS 1775) distalis dorsally tumescent, proximal to apex (Figs 19b, 20, 21c, 23) of vesica, without an apical process; ejacula- Cimex imperialis FABRICIUS 1775: 697 (n.sp.); tory reservoir absent; CAI elongate, claw- FABRICIUS 1781: 339 (description); FABRICIUS shaped, with basally broad, plate-like, with 1787: 280 (description); FABRICIUS 1794: 81 (de- scription); DONOVAN 1805: pl. 3 fig. 2 (habitus) apex narrow, strongly arcuate inwardly; Tetyra imperialis: FABRICIUS 1803: 128 (new com- CAII prominent, bifurcate, CAII(M) basal- bination) ly membraneous, with apex outwardly arcu- Calliphara imperialis: GERMAR 1839: 126 (new ate, heavily sclerotized, with subdistal scle- combination); HERRICH-SCHAEFFER 1840: 83, fig. rotised spine, and moderately sclerotised 529 (description); STÅL 1866: 153 (description); apical bulb-shaped process, CAII(L) thorn- LETHIERRY & SEVERIN 1893: 24 (catalogue); like in lateral view, basally membraneous, SCHOUTEDEN 1904: 32 (list); VAN DUZEE 1905:

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189 (description); KIRKALDY 1909: 297 (cata- connexiva VI-VII with acuminate spines logue); TILLYARD 1926: 149 (diagnosis); MCDON- (Fig. 21b). Male Genitalia: genital opening ALD 1961: 182, figs 29-32 (male genitalia); MC- of pygophore with dorsal and ventral patch- DONALD 1963a: 30 (male genitalia); MCDONALD es of setae; vesica short, with a small subdis- 1963b: 235, figs 17, 18 (female genitalia); KUMAR tal tumescence, ejaculatory apparatus small, 1964: 62 (male genitalia); KUMAR 1965: 53, fig. paired ventral conducting canals, with three 87 (female genitalia); MCDONALD & CASSIS 1984: 566 (description); CASSIS & GROSS 2002: pairs of convolutions; ejaculatory reservoir 589 (catalogue) absent; CAI elongate, S-shaped, apex acute, Callidea imperialis: DALLAS 1851: 24 (new combi- strongly sclerotised; CAII basally membra- nation) neous medially, apex with bifid lobal scle- Diagnosis: Calliphara imperialis is recog- rite; CAIII(D) elongate, narrow, S-shaped, nised by the following characters: dorsum sclerotized process; CAIII(V) absent. Fe- dark orange (Fig. 19b); head and tip of male Terminalia: paratergites VIII large, scutellum iridescent dark-green (Fig. 19b); subtriangular; paratergites IX small, subel- pronotal callosite region with black medial liptoid; gonocoxae I large, subtriangular, spot (Fig. 19b); ventral surface of body flap-like. mostly dark black with iridescent green Measurements. MCDONALD & CASSIS tinge, lateral margins of sternites of pregen- 1984: Table 6. ital abdomen orange (Fig. 21b); dorsum densely punctate (Fig. 19a); abdominal SIV- Material examined: Queensland: 1= 1Y, Blue- VII strongly spinose (Fig. 21b); lateral mar- water Range, N of Townsville, T Woodger & L Y gins of abdominal venter impunctate (Fig. Ring, 29-xii 1990 (AM); 1 , ‘K3613’ (AM); == YY = 21b); CAI elongate, S-shaped; CAII with 3 2 , Brisbane (AMNH); 1 , Palm Is- == YY bifid lobal sclerite; CAIII(D) elongate, nar- land, W Taylor (AMNH); 2 2 , ‘318’ (AMNH). row, S-shaped, sclerotized process; and, Distribution: Calliphara imperialis is en- CAIII(V) absent. demic to Australia (Fig. 23), found primari- Description: Body elongate-ovoid; large, ly in the wet tropics of northeast Queens- males 17-19 mm, females 18-20 mm; land. CASSIS & GROSS (2002) also listed it Colouration. Body bicoloured (Fig. from the Northern Rivers of New South 19b), dorsum mostly burnt-orange with Wales and southeast Queensland, but there head fuscous iridescent green and tip of is some doubt as to whether this species oc- scutellum iridescent green, sometimes cal- curs this far south. We have seen a single losite region of pronotum with large dark specimen from the Northern Territory, medial spot; antennae and labium black; without a specified location. This species legs black, with iridescent green or blue was erroneously recorded from New Zealand tinge; thoracic pleura mostly fuscous, later- (CASSIS & GROSS 2002, references therein). ally with iridescent green hue, metepister- Host plant and biology: No host plant num orange; most of abdominal venter has been recorded. black, with lateral margins orange; termina- lia black with iridescent green hue. Remarks: LYAL (1979) designated a lec- totype, and we have accepted this as the Texture. Dorsum with moderate distri- original description does not indicate the bution of very shallow punctures; abdominal number of specimens in the type series. venter weakly rugulose. LYAL (1979) assigned C. imperialis to the Vestiture. Dorsum glabrous; abdominal Calliphara excellens species-group, on the ba- venter with sparse distribution of decum- sis of the male pygophore and aedeagus. The bent, simple, pale setae. lobal sclerite of CAII is significantly differ- Structure. Antennae: AII(a) very short, ent to all other of the Australian species of ca. 1/3 of AI. Labium: reaching posterior Calliphara, bifid, but not U-shaped as in the margin of abdominal sterna IV. Eyes: not other species. In addition, this species has S- large, about 1/2 length of Head. Pregenital shaped CAI and CAIII(D). Our observa- Abdomen: posterior angles of abdominal tions differ significantly from those of MC-

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DONALD’s (1961) description of the con- Scutellum: mostly burnt orange; thoracic junctival appendages. In addition, MCDON- pleura bicoloured, ventrally orange, lateral- ALD (1961) does not report a setal patch on ly with greenish iridescence; Legs: coxae, the dorsal margin of the pygophore, which is trochanters and proximal 4/5 of femora in contradistinction to LYAL (1979) and our burnt-orange to red; remainder of legs fus- observations a character state present in all cous to bluish-fuscous. Abdominal Venter: species of Calliphara. mostly orange, with broad sublateral green iridescence, with embrowned margins (Fig. Calliphara nobilis (LINNAEUS 1763) (Figs 21c); terminalia embrowned; ventral surface 19c, 21c) Revised Australian Record of pygophore iridescent copper-brown, Cimex nobilis LINNAEUS 1763: 17 (n.sp.); STÅL basally iridescent green (Fig. 21c). 1866: 153 (description) Texture. Body densely punctate, with Cimex pustulatus PANZER 1798: 111 (n.sp.; junior shallow punctures. homonym); LYAL 1979: 172 (synonymy) Scutellera buquetti GUÉRIN-MÉNEVILLE 1838: 159, Vestiture. Abdominal venter with mod- 162 (n.sp.); LYAL 1979: 172 (synonymy) erate distribution of pale, simple setae. Callidea nobilis: GERMAR 1839: 117 (new combi- nation); DALLAS 1851: 25 (list); WALKER 1867: Structure. Eyes: large, about 1/2 length 32 (list); STÅL 1873: 17 (list); ATKINSON 1887: of Head. Antennae: AII(a) a little longer 165 (list); TRYON 1892: 14 (New Guinea); than AI. Labium: reaching between middle LETHIERRY & SEVERIN 1893: 24 (catalogue); DIS- and apex of metacoxae. Male Genitalia: ven- TANT 1902: 53, fig. 23 (description); SCHOUTE- tral margin of male pygophore deeply emar- DEN 1904: 32 (list); KIRKALDY 1909: 298 (cata- ginate; genital opening of pygophore with logue); TAKARA 1957 (host plant); BALOCH, MO- dorsal and ventral patches of setae; vesica HYUDDIN & GHANI 1968 (host plant); HILGEN- short, with arcuate dorsal process; CAII bi- DORF & GOEDEN 1982 (host plant); TOMOKUNI et furcate, CAII(M) mostly membraneous with al. 1993: 218, pl. 89 (description; colour photos); U-shaped lobal sclerite, CAII(L) elongate, KOHNO 2006: 67 (biology; host plants) digitiform, sclerotized, distally recurved; Calliphara buquetti: STÅL 1866: 153 (new combi- CAIII bifurcate, CAIII(D) branch elongate, nation) digitiform, sclerotized; CAIII(V) short, Diagnosis: Calliphara nobilis is recognised mostly membraneous, lobate. Abdominal by the following characters: dorsum irides- Venter: posterior angles of female abdominal cent green; callosite region mostly black SVI-VII with minute acuminate spines; male (Fig. 19c); pronotal black with 8 black spots posterior angles of abdominal sterna bare (Fig. 19c); scutellum with six spots (Fig. (Fig. 21c). Female Terminalia: paratergites 19c); femora orange; abdominal venter VIII large, concave and triangular; parater- mostly orange, with broad sublateral green gites IX smaller, convex and club-shaped, iridescent band (Fig. 21c); eyes large (Fig. fused medially to the tenth sternum; and 19c); lateral margins of abdominal sterna gonocoxae I large, subtriangular, flap-like. densely punctate (Fig. 21c); posterolateral Measurements. BL: 12.89, PW: 8.25, angles of abdominal sterna without spines IOD: 1.98, AI: 0.64, AII(a): 1.04, AII(b) (Fig. 21c); CAII(M) with U-shaped lobal 4.54, AIII: 5.24, AIV: 5.46, LI: 3.61, LII: sclerite CAIII divided; and, vesica short. 3.95, LIII: 3.46, LIV: 2.77. Description: Body moderately large, Specimens examined: Australia: 1=, Northern male 13 mm. Territory, 7 km ESE Smith Point, Colouration. Body mostly green to cop- 11°09’S132°11’E, Cobourg Peninsula, 23 January 1977, ED Edwards (ANIC); Indonesia: 5== 1Y, per-green, with patterned black spots on Sumatra, Dolok Merangir, EW Diehl (AMNH); pronotum (4 spots medially, plus subtriangu- 2YY, East Borneo, Saminedia, Sternitzky coll. lar burnt-orange calli) and scutellum (7 (AMNH); 1=, Sumatra, Stabat, 60 km NW of spots) (Fig. 19c). Antennae: uniformly fus- Medan, 30 m, 23-vi-1974, EW Diehl (AMNH); cous. Pronotum black, tip of scutellum, Philippines: 4== 3YY, Mindanao, Calapan, P hemelytral margins, appendages, connexiva, de Mesa (AMNH). and male and female terminalia black to Distribution: This species is broadly dis- bluish-black, sometimes with a green tinge. tributed in southeast Asia from China (in-

342 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at cluding Taiwan) and Burma in the north, LETHIERRY & SEVERIN 1893: 24 (catalogue); DIS- and through the Indo-Malaysian peninsula, TANT 1899: 38 (synonymy); KIRKALDY 1909: 298 as well as the Philippines and New Guinea. (catalogue); LYAL 1979: 173 (synonymy) Tetrarthria sobria WALKER 1867: 21 (n.sp.); KIRKALDY (1909) tentatively recorded C. nobilis from Australia (Tropical Queens- LETHIERRY & SEVERIN 1893: 25 (catalogue); KIRKALDY 1909: 297 (synonym of Calliphara land?), LYAL (1979) explicitly excluded it. praslinia); LYAL 1979: 173 (synonymy) The specimen from the Northern Territory Callidea erythrospila WALKER 1867: 33 (n.sp.); (Fig. 23) is the first precise Australia record. LETHIERRY & SEVERIN 1893: 23 (as Calliphara bil- Host plants and biology: There was no lardierii ?ssp.); DISTANT 1899: 38 (synonymy); host plant record for the Northern Territory KIRKALDY 1909: 298 (catalogue; synonymy); SCHOUTEDEN 1904: 32 (synonymy) specimen listed above. There are five host Callidea semirufa WALKER 1867: 34 (n.sp.); plants recorded for this species from outside LETHIERRY & SEVERIN 1893: 23 (as Calliphara bil- of Australia (Table 1). lardierii ?ssp.); DISTANT 1899: 38 (synonymy); Remarks: In LYAL’s (1979) classification, KIRKALDY 1909: 297 (catalogue; synonymy); DIS- C. nobilis belongs to the Calliphara excellens TANT 1899: 38 (synonymy); SCHOUTEDEN 1904: 32 (synonymy) species group, the latter established primari- Callidea biplaga WALKER 1867: 35 (n.sp.); ly on the condition of the conjunctival ap- LETHIERRY & SEVERIN 1893: 23 (as Calliphara bil- pendages. This species group comprises sev- lardierii ?ssp.); DISTANT 1899: 38 (synonymy); en species, including two Australian species KIRKALDY 1909: 297 (catalogue; synonymy); (C. imperialis and C. regalis) additional to C. SCHOUTEDEN 1904: 32 (synonymy) nobilis, as well as three species from Indone- Diagnosis: Calliphara regalis is recognised sia, C. lanceolata, C. regia and C. excellens, by the following characters: dorsum orange, with the last also occurring in the Philip- with head and tip of scutellum iridescent pines. The relationships of these taxa have dark-green (Figs 1a, 19d); base of femora or- not been analysed, although, C. nobilis ange, remainder orange-fuscous; pregenital shares numerous similarities in the male abdominal venter mostly orange, sometimes genitalia with C. regalis, including a short with trichobothrial area dusty; abdominal vesica, simple ejaculatory apparatus, and SIV-VII minutely spinose; CAI(D) lobate very similar conjunctival appendages. They (Figs 22c, d); CAI(V) elongate, outwardly are easily separated externally by coloura- arcuate (Figs 22c, d); CAII(M) with distal tion patterns alone (cf. Figs 19a-d). bifid lobal sclerite (Figs 22c, d); CAIII elon-

Calliphara regalis (FABRICIUS 1775) gate, digitiform (Figs 22c, d); and, vesica (Figs 1a, 19d, 20, 22, 23) short, with apical hood (Figs 21c, d). Cimex regalis FABRICIUS 1775: 697 (n.sp.); FABRI- Description: Body large, males 16-17 CIUS 1781: 339 (description); FABRICIUS 1794: 80 mm, females 17-19 mm. (description); DONOVAN 1805: pl. 3 fig. 3 (habi- tus) Colouration. (Australian morph) Body Tetyra regalis: FABRICIUS 1803: 28 (new combina- bicoloured (Figs 1a, 19d); dorsum mostly tion); GERMAR 1839: 127 (description) golden-orange with head fuscous iridescent Calliphara regalis: GERMAR 1839: 127 (new com- green and tip of scutellum with a round fus- bination); STÅL 1866: 153 (description); cous subdistal spot; antennae black; labium LETHIERRY & SEVERIN 1893: 24 (catalogue); DIS- mostly orange-black, LIV fuscous. Legs: cox- TANT 1899: 38 (list; synonymy); VAN DUZEE ae, trochanters and proximal 4/5 of femora 1905: 189 (diagnosis); SCHOUTEDEN 1904: 32 orange-red, apex of femora, tibiae and tarsi (list); KIRKALDY 1909: 297 (catalogue); SCHOUT- orange-fuscous. Thoracic pleura mostly EDEN 1933: 46 (Indonesia); MCDONALD 1961: golden-orange, with irregular fuscous mark- 182, figs 33-38 (male genitalia); MCDONALD ings. Pregenital Abdomen: venter mostly or- 1963a: 30 (male genitalia); BLACK 1968: 573 ange, with SII/III boundary and areas (Melanesia); MCDONALD & CASSIS 1984: 563 bounding trichobothria fuscous. Female Ter- (description); CASSIS & GROSS 2002: 589 (cata- logue) minalia: with fuscous highlighting. Male Caliphara (Chrysocoris) imperialis: STÅL 1873: 18 Genitalia: pygophore fuscous-iridiscent (subgeneric arrangement) green; with lateral margins orange; termina- Callidea eximia VOLLENHOVEN 1863: 20 (n.sp.); lia black with iridescent green hue.

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Texture. Pronotum and scutellum with moderate distribution of fine punctures, less densely distributed on callosite region and anterior callus of scutellum; propleuron and metepimeron with sparse distribution of deep punctures; abdominal venter weakly rugulose, more rugopunctate laterally. Vestiture. Dorsum glabrous; abdominal venter with sparse distribution of pale, semi- erect, pale setae. Structure. Head: eyes large, greater than 1/3 length of Head. Antennae: AII(a) short, little more than 1/2 of AI; Labium: reaching posterior margin of metacoxae to abdominal sterna II. Pregenital Abdomen: posterior angles of SIV-VII minutely spin- ose. Male Genitalia: ventral margin of male pygophore rounded, genital opening of py- gophore with dorsal and ventral patches of setae (Figs 20f, 22a); parameres with small medial tumescence, setose (Fig. 22b); ejacu- latory apparatus small, ventral conducting canal short, with two pairs of convolutions (Figs 22c, d); ejaculatory reservoir short, heavily sclerotized (Figs 22c, d); CAI(D) lo- bate, weakly sclerotized medially, CAI(V) elongate, outwardly arcuate, heavily sclero- tized (Figs 22c, d); CAII(L) with large, membraneous lobe; CAII(M) with distal bi- fid lobal sclerite (Figs 22c, d); CAIII greatly elongate, digitiform lobal sclerite (Figs 22c, d); vesica short, secondary gonopore posi- tioned just post-thecal margin, with a great- ly enlarged, hooded apical process (Fig. 22d). Female Terminalia: paratergites VIII large, subtriangular, medially separated; paratergites IX smaller, subelliptoid; gono- coxae I large, subtriangular, flap-like. Sper- matheca: reservoir elliptoid.

Measurements. MCDONALD & CASSIS 1984: Table 6.

Type material examined: Callidea biplaga WALKER: Holotype, =, ‘Aru Isl.’, 58-48’, ’42 Callidea bipla- gia’ ‘B.M. Hem. Type No. 471’ (BMNH; der- mestid damage; without lectotype label); Callidea semirufa WALKER: Holotype, Y, ‘Waigou’, ‘Saun- ders 65-13’, ‘Callidea semirufa WALK. (type) ’, Fig. 24: Habitus of Australian Scutellerinae species (a) Calliscyta stalii, Y (b) Heissiphara ‘B.M. Hem. Type No. 472’ (BMNH; dermestid minuta nov.sp., Y (c) Choerocoris lattini nov.sp., Y (d) Choerocoris paganus, = (e) damage; without lectotype label); Callidea ery- Choerocoris grossi nov.sp., = (f) Choerocoris variegatus, =. Scale bar = 1 mm. throspila WALKER: Lectotype, =, ‘Cer.’, ‘Saunders 65-13’, ‘Callidea erythrospila’, ‘B.M. Hem. Type No. 473’ (BMNH; damaged; without lectotype label); Tetrarthria sobria WALKER: Holotype, Y,

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‘New Heb’, ‘66-12’, ‘Tetrarthria sobria WALK’, Calliscyta STÅL 1873 (Figs 21d, 24a, 25, ‘B.M. Hem. Type No. 468’ (BMNH; damaged; 26, 27) Revised Status without lectotype label). Calliscyta STÅL 1873: 9 (gen.n.); LETHIERRY & Other material examined: Queensland: 1=, SEVERIN 1893: 31 (catalogue); SCHOUTEDEN Cairns, July 1954, A Homes (AM); 1Y, Etty Bay, 1904: 26 (description); KIRKALDY 1909: 301 (cat- 8 km SE Innisfail, 13-viii-1983, NW Rodd (AM); alogue); MCDONALD & CASSIS 1984: 538 (junior = 1 , Clump Point, 6 March 1964, IFB Common synonym of Choerocoris); CASSIS & GROSS 2002: Y & MS Upton (ANIC); 1 , Lloyd Bay, 3 miles N 592 (junior synonym of Choerocoris) Claudie River, 14-i-1972, DK McAlpine & GA Holloway (AM); 1=, Palm Island (AMNH); Diagnosis: Calliscyta is recognised by the 2== 1Y, ‘318’ (AMNH). following combination of characters: elon- Distribution: Calliphara regalis is wide- gate-ovoid (Fig. 24a); head strongly de- spread outside Australia from Indonesia (as flexed (Fig. 24c); jugal margins rounded far west as Sumatra) to New Caledonia and (Fig. 24c); anterolateral margins of prono- Vanuatu in the east. In Australia, it is tum strongly divergent, rectilinear (Figs known from the Torres Strait Islands to the 24a, 25a), carinate in profile (Figs 25c, d); wet tropics of Queensland (Fig. 23). anterolateral margins of scutellum foveate (Fig. 25a); peritreme broad, subtriangular Host plant and biology: No host plant (Fig. 25d); evaporative areas reduced (Fig. has been recorded for this species. This 24d); posterior margin of pygophore emar- species has been found in aggregations on ginate (Fig. 25e, f); ejaculatory apparatus leaves (Fig. 1a). prominent, convoluted ventral conducting

Remarks: LYAL (1979) designated all of canal present (Figs 26c, d); CAI absent (Fig. the Walker types of the junior synonyms as 26c); CAII symmetrically bifurcate (Figs lectotypes. The lectotype designations were 26c, d); CAIII fused medially (Figs 26c, d); unnecessary as the original descriptions re- and, vesica elongate (Figs 26c, d). fer to a single specimen, and only a single Type species: Calidea stalii VOLLENHOVEN 1863, specimen of each available name was found monotypy in the BMNH. In addition, we found that Description: Moderately sized; elongate- he did not add lectotype labels to the speci- ovoid (Fig. 24a); dorsum mostly dark with mens. We formally reject these lectotype orange to red markings (Fig. 24a); dorsally designations and regard them as holotypes. and ventrally strongly convex (Fig. 25c); body mostly punctulate (Fig. 24a); tip of LYAL (1979) placed C. regalis in the Cal- wing visible beyond emarginate apex of liphara excellens species-group on the basis of Scutellum. Head: transverse; subtriangular the male pygophoral and aedeagal charac- (Fig. 25a), moderately deflexed; lateral mar- ters. MCDONALD (1961) described its male gins of jugae rounded, punctulate (Fig. 25a); genitalia in detail, showing the elongate, bucculae narrow. Antennae: AI, AII(a) and digitiform CAI(V) and CAIII. Our dissec- AII(b) short, AIV longest segment; AII(b)- tions from across the range of this species AIV compressed. Labium: reaching base of confirm McDonald’s observations. It ex- metacoxae. Pronotum: broadly trapezoidal hibits remarkable colour variation; in Aus- (Fig. 24a), strongly convex; anterolateral tralia, all the specimens we observed have margins elongate, rectilinear, carinate; the dorsum mostly orange, with the head iri- humeral angles rounded; posterolateral mar- descent green and the subapex of the scutel- gins short, rounded; posterior margin recti- lum with a subdistal iridescent fuscous- linear. Scutellum: elongate (Fig. 24a), green; sometimes the pronotum is darker. In strongly convex in profile, anterolateral an- Melanesia, this species is extremely vari- gles depressed; apex truncate (Fig. 24a); able, but rarely as in the Australian colour forewings weakly exposed basally and cau- morph, and can be mostly iridescent blue or dally. Thoracic sterna: without prominent green, or copper-green, with up to eight carination (Fig. 25b). Thoracic pleura: ante- black spots on the scutellum. The male rior margin of proepisternum lobate, moder- aedeagi of all the colour morphs are identi- ately explanate (Fig. 25b). Thoracic Pleura: cal, and the colour variation is extreme if external efferent system of metathoracic not continuous. glands moderately developed (Fig. 25d), os-

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Fig. 25: Scanning electron micrographs of tiole small; peritremal plate large, strongly margin with deep v-shaped notch (Figs 25e, key characters of Calliscyta stalii (a) Head, convex (Fig 25b), without prominent medi- f, 26a), with moderate distribution of setae dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External al sulcation; evaporative areas restricted to (Fig. 25f); parameres basally columnar with efferent system of metathoracic glands, either side of anterior and posterior margins shallow, hook-shaped apex, tip rounded ventral view (e) male pygophore, ventral of the peritremal plate, not extending to (Fig. 26b); aedeagus with heavily-sclerotized view (f) male pygophore, dorsal view. mesepimeron (Fig. 25d). Male abdomen: box-shaped phallotheca, without processes Abbreviations: Ea = evaporative area; Go = genital opening; P = pronotum; P(al) = strongly convex, broad; abdominal tri- (Figs 26c, d); two conjunctival appendages, anterolateral margin of pronotum; Pe = chobothria in recessed area; urite VIII hid- CAII pair symmetrical membraneous, bifid, peritreme; Pe(k) = keel of anterior margin den. Male Genitalia: pygophore moderately CAII(M) and CAII(L) with apical conical of proepisternum; P(r) = right paramere; Pr sized, lozenge-shaped, ventral surface sinu- sclerites (Figs 26c, d); CAIII bifid, large, = proctiger; EES = external efferent system. ate in profile, medially excavate, posterior heavily-sclerotized, U-shaped, medially

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Fig. 26: Male genitalia of Calliscyta stalii (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) apex of aedeagus, ventral view. Abbreviations: C = crown of paramere; CAI(V) = ventral branch of first conjunctival appendage; CAI(D) = dorsal branch of first conjunctival appendage; CAII(L) = lateral branch of second conjunctival appendage; CAII(M) = medial branch of second conjunctival appendage; CAIII = third conjunctival appendage; DCC = dorsal conducting canal; DS(D) = ductus seminis distalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; SP = setose patches of male genital opening; VCC = ventral conducting canal; V = vesica; VM = ventral margin of genital opening.

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In contrast, the conjunctival appendages of C. stalii and C. paganus are more alike, than the latter is with its congener C. variegatus. MCDONALD (1963a) reported on the signif- icance of the bifid condition of the CAII and CAIII conjunctival appendages in these taxa. However, our observation reveal that these character states are broadly held in scutellerids. We consider the shape of the conjunctiva as of importance in separating species, but are doubtful indicators of phylo- genetic significance. MCDONALD & CASSIS (1984) also indicated that the female geni- talia of Calliscyta and Choerocoris are closely aligned, but our observations indicate no special similarities in the external female genitalia and spermatheca. The conditions described in MCDONALD (1963b) are more descriptive of the Scutellerinae, rather than at any subordinate taxonomic level. In contrast, Calliscyta stalii, differs from Choerocoris, by the following characters: lat- Fig. 27: Distribution of Scutellerinae fused at base (Figs 26c,d); ductus seminis eral margins of jugae rounded (cf. carinate), species (a) Calliscyta stalii (b) Cantao proximalis narrow; ejaculatory apparatus forewings extending beyond apex of scutel- parentum (c) Heissiphara minuta nov.sp. with heavily-sclerotized ventral convoluted lum (cf. covered), peritreme and evaporative conducting canals (Figs 26c, d); ejaculatory areas of the metathoracic glands greatly re- reservoir heavily-sclerotized, elongate-ovoid duced (cf. extensive), ventral margin of the (Figs 26c, d): dorsal conducting canal broad, male pygophore emarginate (cf. entire), and strongly sclerotized (Figs 26c, d); ductus tapered ductus seminis distalis. These char- seminis distalis strongly sclerotized (Figs acter differences are not only sufficient in 26c, d); vesica broad (Figs 26c, d). Female recognising Calliscyta, but also provide sig- Terminalia: paratergites VIII broad, large, nificant differentiation from Choerocoris, medially emarginate; paratergites IX small, such that the closeness of the relationship of subelliptoid; gonocoxae I subtriangular, these two genera is in question. with posterior margins concave; fecunda- tion canal short; spermathecal reservoir cir- Calliscyta stalii (VOLLENHOVEN 1863) cular, large, strongly thickened; fecundation (Figs 21d, 24a, 25, 26, 27) canal short, narrow spermathecal bulb heav- Revised Status ily sclerotized, with proximal and distal Calidea stalii VOLLENHOVEN 1863: 24 (n.sp.) flanges. Calliscyta stalii: STÅL 1873: 24 (new combina- tion); LETHIERRY & SEVERIN 1893: 31 (cata- Diversity and distribution: Calliscyta is a logue); SCHOUTEDEN 1904: 25 (list, habitus); monotypic genus, restricted to tropical east- VAN DUZEE 1905: 190 (distribution); BERGROTH ern Queensland. 1908: 296 (list); KIRKALDY 1909: 301 (list); WU 1933: 229 (China, incorrect record) Included species: Choerocoris stalii: MCDONALD &CASSIS 1984: C. stalii (VOLLENHOVEN 1863) Australia 561 (new combination); CASSIS &GROSS 2002: 594 (catalogue) Remarks: MCDONALD & CASSIS (1984) Calliscyta australis DISTANT 1899: 40 (n.sp.); synonymized Calliscyta with Choerocoris LETHIERRY & SEVERIN 1893: 31 (catalogue); based on similarities of the male and female SCHOUTEDEN 1904: 25 (list); BERGROTH 1908: genitalia. Males of Calliscyta stalii differ sig- 296 (list); KIRKALDY 1909: 301 (list); MCDONALD nificantly from Choerocoris paganus and C. 1963a: 26 (male genitalia); MCDONALD &CAS- variegatus, by having the ductus seminis dis- SIS 1984: 561 (synonymy); CASSIS &GROSS talis tapered as opposed to greatly incrassate. 2002: 594 (catalogue; synonymy)

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Diagnosis: Calliscyta stalii is recognised spots (Fig. 21d); and, genitalia as in generic Fig. 28: Habitus of Australian Scutellerinae = by the following combination of characters: diagnosis (Figs 26a-d). (a) Cantao parentum, (b) Scutiphora pedicellata, Y (c) Lampromicra aerea, = dorsum mostly red-fuscous, with orange = Description: Body moderately large, (d) Lampromicra regia, (e) Lampromicra markings on pronotum and scutellum (Fig. senator, =. Scale bars = 1mm. males 13-14.7 mm, females 13.6-14.7 mm; 24a); AIV longest segment; labium reaching dorsum strongly convex. midpoint of metacoxae; anterolateral mar- gins of pronotum elongate; ventral surface Colouration. Dorsum mostly red-fus- of legs with thick orange setae; abdominal cous and green iridescence, with orange venter mostly red with sublateral fuscous markings on pronotum and scutellum (Fig.

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24a). Head: mostly fuscous, sometimes with specified locality in Timor. DISTANT (1899) green iridescence, jugae orange-red. Anten- subsequently described an additional species, nae: uniformly fuscous. Labium: fuscous. C. australis, from Queensland. MCDONALD Pronotum: mostly red-fuscous, callosite re- & CASSIS (1984) synonymized the latter on gion and pronotal midline most often with the basis of similarities of the male genitalia. yellow-red to orange-red marking. Scutel- Calliscyta stalii is not common in collec- lum: mostly red-fuscous, anterolateral re- tions and often has a greasy appearance. It is gions with yellow-red to orange-red mark- much larger than species of Choerocoris, and ings. Thoracic Pleura: mostly yellow to or- more elongate-ovoid, than ovoid. Re-exam- ange with fuscous punctations. Legs: uni- ination of the type material of C. stalii and formly fuscous, often with green iridescence. C. australis confirm the synonymy given by Pregenital Abdomen: mostly red, with fus- MCDONALD & CASSIS (1984). This species cous patches bounding spiracular + tri- shows less variation in size and colouration chobothria region (Fig. 21d). than in most other scutellerine species. Texture. Dorsum densely punctate, with There is some intraspecific variation in the moderately deep punctures; callosite region colour patterning of the pronotum and of pronotum partly impunctate; underside of scutellum, particularly in the extent of the head and thoracic pleura densely punctate; lighter markings. abdominal venter weakly rugopunctate. Cantao AMYOT & SERVILLE 1843 Vestiture. Dorsum glabrous; ventral sur- (Figs 1b, 21e, 27, 28a, 29, 30) faces of legs setose, with thick yellow setae; Cantao AMYOT & SERVILLE 1843: 29 (gen.n.); abdominal venter with a few scattered setae. DALLAS 1851: 3, 17 (key, list); STÅL 1865: 33 Structure. Antennae: relatively short; (key); MAYR 1866: 14 (description); STÅL 1873: AIV longest segment, little longer than AI- 10 (list); LETHIERRY &SEVERIN 1893: 18 (cata- II; AII(a) shortest segment; AI & AII(b) logue); DISTANT 1902: 42 (description); SCHOUT- EDEN 1904: 18 (description); KIRKALDY 1909: 307 subequal in length; AIII and AIV weakly (catalogue); MCDONALD & CASSIS 1984: 550, flattened. Labium: flattened, reaching mid- 552 (key, description); CASSIS & GROSS 2002: point of metacoxae; LII longest segment; LI 591 (catalogue) shortest segment; LIII-LIV subequal in Cantao (Iostethus) STÅL 1873: 10 (gen.n.); length. Pronotum: anterolateral margins KIRKALDY 1909: 307 (catalogue) greatly elongate, more than 2x length of Type species: Cantao: Cimex ocellatus THUNBERG posterolateral margins. Pregenital Ab- 1789, subsequent designation, KIRKALDY 1909: 307; domen: connexival margins entire, without Cantao (Iostethus): Calidea parentum WHITE 1839: posterolateral spines or nodules (Fig. 21d). 85, subsequent designation, KIRKALDY 1909: 307 Genitalia: as in generic description. Diagnosis: Cantao is recognised by the

Measurements. MCDONALD &CASSIS following characters: elongate-ovoid (Figs 1984: Table 5 (as Choerocoris stalii) 1b, 28a); large size; bicoloured, orange or red with fuscous to dark blue markings (Figs 1b, Other material examined: Queensland: 2== 28a), sometimes with green iridescence ven- YY 2 , South Percy Island, NW Bay, 23-29 No- trally; body smooth, punctulate to rugo- vember 1992, GB Monteith, G Thompson, D punctulate (Figs 1b, 28a); hemelytra ex- Cook & H Janetzki (QM). tending beyond tip of abdomen; lateral mar- Distribution: Calliscyta stalii is known gins of jugae almost straight (Fig. 29a), car- from North Queensland (Fig. 27). inate in profile (Fig. 29b); anterolateral Host records and biology: Monteith margins of pronotum carinate; anterolateral (pers. comm.) found large numbers of Callis- angles of scutellum foveate (Figs 1b, 28a); cyta stalii in the leaf axils of a species of the apex of scutellum truncate (Fig. 28a); per- monocot genus Pandanus (Pandancaceae). itreme greatly enlarged, subtriangular, raised, sulcate at base (Fig. 29d); abdominal It is uncertain whether this is a food plant or SVIII visible (Fig. 29e); male pygophore a site. large, ventrally oriented (Fig. 29e); ejacula- Remarks: VOLLENHOVEN (1863) origi- tory apparatus with convoluted ventral con- nally described Calliscyta stalii from an un- ducting canals (Figs 30c, d); CAII undivid-

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Fig. 29: Scanning electron micrographs of key characters of Cantao parentum (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ea = evaporative area; Go = genital opening; P = pronotum; P(al) = anterolateral margin of pronotum; Pe = peritreme; Pe(k) = keel of anterior margin of proepisternum; P(r) = right paramere; Pr = proctiger; EES = external efferent system. Scale bars = 1 mm.

ed, with blunt lobal sclerite (Figs 30c, d); areas posteriad to peritreme. Head: trans- spermathecal reservoir large, oval; sper- verse, convex in dorsal view; jugal margins mathecal pump with proximal and distal carinate; jugae rugulose; bucculae narrow, flanges. sometimes margins weakly excavate. An- Description: Body elongate-ovoid, tennae: AI and AII(a) short, AII(b)-AIV hemelytra extending beyond truncate tip of subequal in length. Labium: reaching be- scutellum (Figs 1b, 28a); dorsum ground tween apices of metacoxae and basal ab- colour yellow, orange or red-brown with pat- dominal sterna. Pronotum: broadly trape- terned fuscous to black markings (Figs 1b, zoidal, wider than long (Figs 1b, 28a), even- 28a), venter with green-fuscous iridescent ly convex in profile, without a transverse patterned markings; body smooth, punctu- furrow, posteriad to callosite region; anterior late to rugo-punctulate; ventral surface margin deeply excavate (Figs 1b, 28a), densely setate with rows of setae on anterior sometimes weakly depressed; anterolateral margin of proepisternum and evaporative margins strongly carinate, weakly to strong-

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Fig. 30: Male genitalia of Cantao parentum (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) apex of aedeagus, ventral view. Abbreviations: C = crown of paramere; CAII = second conjunctival appendage; CAIII = third conjunctival appendage; DCC = dorsal conducting canal; DS(D) = ductus seminis distalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; VCC = ventral conducting canal; V = vesica; VM = ventral margin of genital opening.

352 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at ly concave (Figs 1b, 28a); posterior margin & STONEDAHL 1986). The Australian weakly excavate (Fig. 28a); humeral angles species C. parentum is found through rounded (Figs 1b, 28a) to prominently spin- coastal Queensland, as far south as the ose; posterior angles, with posteriorly pro- Northern Rivers district of New South jecting thorn-like process. Scutellum: elon- Wales. There is some doubt as to the native gate (Fig. 28a), strongly convex in profile; distribution and identity of C. africanus. anterolateral region deeply foveate (Figs 1b, HORVÁTH’S (1892) original description 28a), forewings minimally exposed basally; gives the Congo as the type locality. truncate apically, wings visible (Figs 1b, SCHOUTEDEN (1904) suggested that the 28a). Thoracic sterna: without carination species could be either from the Congo or (Fig. 29c). Thoracic pleura: anterior margin Queensland. MCDONALD (1988) queried if of proepisternum thickened (Fig. 29c). Tho- C. africanus is a synonym of C. ocellatus, racic Pleura: ostiole small, with greatly en- based on identified material of the latter larged subtriangular peritreme, with shallow from Zaire. Cantao ocellatus is the most basal sulcation (Fig. 29d); evaporative areas broadly distributed species in the genus, restricted to anterior and posterior margins and is found from the Afrotropical region to of peritreme, not extending laterally beyond Papua New Guinea, as well as many coun- peritreme (Fig. 29d); mesepimeron without tries in the Oriental region, and extending evaporative areas. Male Pregenital Ab- westward as far as . The other three domen: moderately convex, broad; abdomi- species of Cantao exhibit more restricted nal trichobothria in recessed area; SVIII vis- distributions in the eastern Oriental region ible, caudally extending well beyond base of and the Australian region. pygophore (Fig. 29e). Male Genitalia: py- Included species: gophore large, barrel-shaped, ventral surface C. africanus HORVÁTH 1892 sometimes with excavation, ventral margin Afrotropical region sometimes with a U- or V-shaped projection C. ocellatus (THUNBERG 1784) medially (Figs 29e, f); genital opening broad Afrotropical (Zaire), (Fig. 30a), suboval, sometimes with ventral Oriental (broadly) and bifid pygophoral process; parameres hook- Palearctic (Regions) regions, shaped, with apices rounded (Fig. 30b) to Papua New Guinea acute; aedeagus with moderately sclerotized C. variabilis (MONTROUZIER 1855) phallotheca, subconical, without processes Indonesia, Philippines, (Figs 30a, b); CAII membraneous, with Papua New Guinea blunt lobal sclerite, basally with area of scle- C. parentum (WHITE 1839) Australia rotisation (Figs 30c, d); CAIII U-shaped, C. purpuratus (WESTWOOD 1837) heavily sclerotized (Fig. 30d); ductus semin- Indonesia (Banda, Timor) is proximalis narrow, within a phallothecal extension (Fig. 30c); ejaculatory apparatus Remarks: SCHOUTEDEN (1904) divided with highly convoluted ventral conducting Cantao into two subgenera; Cantao (Cantao) canal, with up to 12 convolutions (Figs 30c, and Cantao (Iostethus), based on the pres- d); ejaculatory reservoir sclerotised (Fig. ence or absence of punctures, respectively. 30c); ductus seminis dorsalis and vesica with MCDONALD (1988) synonymized these two subapical tumescence (Fig. 30c). Female subgenera, implicitly dismissing the impor- Terminalia: caudally oriented, sometimes tance of punctation. We endorse this view, recessed; paratergites VIII large, subtriangu- particularly as the supposedly smooth body lar; paratergites IX small to absent. Sper- of C. parentum and C. purpuratus has minor matheca: base of spermatheca strongly scle- punctulation. The male and female geni- rotized; fecundation canal elongate, moder- talia of all Cantao species are alike. In males, ately sclerotized; spermathecal reservoir the aedeagus has two pairs of conjunctival oval, robust; sclerotized proximal and distal appendages (CAII and CAIII), an ejacula- flanges present; and, usually, with heavily tory apparatus with a ventral convoluted sclerotized oval bulb. conducting canal; typical of scutellerines. In Diversity and distribution: Cantao has addition, the male abdominal SVIII is visi- an Indo-Pacific distribution (sensu SCHUH ble, overlapping the basal margin of the Py-

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gophore. The females have terminally ori- with lateral margins weakly serrate (Fig. ented external genitalia, and the spermath- 30d); female terminalia strongly recessed; ecal reservoir is oval and large. In Cantao and, paratergites IX present; there is surprising variation in the develop- Description: Body elongate-ovoid; large, ment of paratergites IX, which can be pres- males9.2-22.5 mm, females 15.5-24.6 mm. ent (parentum) or reduced/absent (africanus, ocellatus and variabilis). Colouration. Dorsum mostly orange, with patterened black spots (Figs 1b, 28a); The sister-genus of Cantao is not appar- body mostly iridescent green-black ventrally. ent, as it shares a number of features with Head: medial regions of clypeus and frons, numerous Old World genera. MCDONALD and vertex green-black, often with iridescent (1988) suggested that Cantao is closely re- green tinge; jugae and frons orange; antennae lated to Calliphara, Callidea LAPORTE and and labium fuscous, often with reddish tinge; Chrysocoris, but gave no morphological basis underside of head red-fuscous; antennae and for such an hypothesis. Cantao, like other labium uniformly fuscous. Pronotum: mostly scutellerine genera such as Chrysophara, orange, with a pair of small black spots, sub- Lampromicra and Scutiphora, has the fore- laterally at base of callosite region (Figs 1b, wings extending beyond the tip of the 28a). Scutellum: mostly orange, with eight scutellum and the female external genitalia black spots, three pair sublaterally from ante- terminal in orientation. However, the exter- rior margin to just beyond midlength, and nal efferent system of the metathoracic single spot along midline at anterior margin glands of Cantao is distinctive, with the and single spot submedially (Figs 1b, 28a). evaporative areas reduced and the peritreme Thoracic pleura: mostly fuscous with purple greatly enlarged. This is unlike that found in to green iridescence, with posterior region of genera such as Calliphara, Chrysocoris, Lam- proepimeron orange. Legs: mostly fuscous, of- promicra and Scutiphora, and similar to the ten with purple to green iridescence, coxae condition found in Calliscyta. with orange shading. Pregenital Abdomen: venter mostly orange, SII uniformly orange, Cantao parentum (WHITE 1839) SIII-SVI with lateral, suboval fuscous mark- (Figs 1b, 21e, 27, 28a, 29, 30) ings (Fig. 21e); SVII medially with large iri- Calidea parentum WHITE 1839: 542 (n.sp.); descent green-fuscous markings, trichoboth- WALKER 1867: 14 (list); LETHIERRY & SEVERIN rial region with short black spot (Fig. 21e); 1893: 18 (catalogue); SCHOUTEDEN 1904: 19 male SVIII mostly orange with iridescent (list); KIRKALDY 1909: 308 (catalogue); TILLYARD green tinge (Fig. 21e). Female Terminalia: or- 1926: 149 (description); MCDONALD 1961: 174, 185, figs 1-4 (male genitalia); MCDONALD 1963a: ange. Male Pygophore: iridscent dark green. 30 (male genitalia); MCDONALD 1963b: 233, figs Texture. Dorsum smooth to weakly 10-12 (female genitalia); MCDONALD 1963c 285 punctulate. (biology); KUMAR 1964: 58 (morphology); KU- Vestiture. MAR 1965: 41 (male genitalia); MCDONALD & Body with moderate distribu- CASSIS 1984: 552 (description); MCDONALD tion of simple, short pale setae; more dense- 1988: 293 (key, synonymy); CASSIS & GROSS ly distributed on head, thoracic pleura and 2002: 592 (catalogue); MILLAR 2005: 78 (glandu- abdominal venter. lar chemistry) Structure. Antennae: AI shortest seg- Cantao pulcher SCHOUTEDEN 1906: 137 (n.sp.); ment; AII(b)- AIV roughly subequal, AIV MCDONALD & CASSIS 1984: 522 (synonymy) longest segment. Labium: reaching between Diagnosis: Cantao parentum is recognised apex of metacoxae and posterior margin of by the following combination of characters: abdominal SII; LI shortest segment; LI dorsum orange, pronotum with pair of black longest segment; LIII & LIV subequal. spots, and scutellum with eight black spots Pronotum: humeral angles rounded (Figs 1b, (Figs 1b, 28a); humeral angles rounded (Figs 28a). Pregenital Abdomen: male abdominal 1b, 28a); male pygophore with ventral SVIII visible, overlapping posterior margin forked process (Figs 29e, f); CAII with basal of Pygophore. Male Genitalia: genital open- sclerotisation and apical, blunt lobal sclerite ing of male pygophore with ventral forked (Fig. 30c, d); CAIII fused basally, U-shaped, process, branches divergent, dorsolaterally

354 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at strongly tumose (Figs 29e, f, 30a); parameres ond Australian species of Cantao, C. pulcher, large, crown weakly hooked, weakly flat- from a male specimen from Queensland. tened, with prominent basal flange (Fig. MCDONALD & CASSIS (1984) synonymized 30b); ejaculatory apparatus well-developed, the latter with C. parentum, regarding it, at heavily sclerotized; ventral conducting most, as a colour variation. Our observation canal elongate, with 10-12 convolutions indicate that there are variations in the (Figs 30c, d); CAII singular, membraneous colour patterning of this species, yet there is lobe, outer margin at base sclerotized, apex almost no variation in the male genitalia with truncate lobal sclerite (Figs 30c, d); and we thus endorse this synonymy. CAIII U-shaped, convergent apically, fused post-thecal margin, heavily sclerotized, mar- Cantao parentum is a distinctive compo- gins weakly serrate (Figs 30c, d); vesica nent of the Australia jewel bug fauna. Some elongate, subdistally swollen (Figs 30c, d). individuals of both sexes are over 20 mm in Female Terminalia: caudal in orientation, length. The body is also relatively less con- recessed; paratergites VIII subtriangular, de- vex than all the other species of jewel bugs, pressed medially; paratergites IX small, sub- and has a bright orange dorsum, with black oval, gonocoxae I large, subelliptoid, ta- spots (Fig. 28a). The male genitalia is also pered medially. Spermatheca: proximal fe- distinctive in that CAII is not subdivided cundation canal elongate, heavily sclero- (Figs 30c, d). It is easily differentiated from tized; reservoir relatively small, oval, heavi- the other species of Cantao (all of which are ly sclerotized; distal fecundation canal mod- found outside of Australia), by the arrange- erately elongate; proximal and distal flanges ment of black spots on the dorsum. In this present; bulb oval, heavily sclerotized. character, it resembles C. ocellatus, but is

Measurements. MCDONALD & CASSIS distinguished from the latter by having 1984: Table 3. rounded humeral angles (cf. spinose angles) and differences in the male and female gen- Material examined: Queensland: 2== 1Y, italia. MCDONALD (1988) provides a key to Lockerbie Scrub, Cape York, 11 April 1975, MS Moulds (AM); 2YY, North Pine River, A Mus- species of Cantao. grave, 30 December 1926 (AM; 58634); 1Y, Surfers Paradise, 10 August 1953 (AM); New Choerocoris DALLAS 1851 (Figs 24c-f, South Wales: 2==, Lismore, A Musgrave, 1 Jan- 31, 32, 33, 34, 35, 36, Table 7) uary 1929 (AM; K47241); 1Y, Tweed Rivers, C Choerocoris DALLAS 1851: 29 (gen. nov.); MAYR Gibbons (AM; K49069). 1866: 22 (description); STÅL 1865: 33 (key); STÅL 1873: 13 (description); LETHIERRY & SEV- Distribution: This species is distributed ERIN 1893: 21 (catalogue); SCHOUTEDEN 1904: 38 in tropical areas of Queensland and the sub- (description); KIRKALDY 1909: 291 (catalogue); tropical areas of eastern Queensland and GROSS 1975: 93 (description); MCDONALD & the Northern Rivers of New South Wales CASSIS 1984: 558 (description); CASSIS & GROSS (Fig. 27). 2002: 592 (catalogue) Host plants and biology: MCDONALD Type species: Cimex paganus FABRICIUS 1775, sub- (1963c) described the biology of this sequent designation: SCHOUTEDEN 1904: 28 species, including life history attributes and Diagnosis: Choerocoris is recognised by host plant associations. Cantao parentum is the following characters: elongate-ovoid (Figs known to breed on three species of the eu- 24c-f); bicoloured, orange or red with fuscous phorb genus Mallotus; M. claoxyloides, M. to dark blue markings, sometimes with green discolor and M. philippensis. A single speci- iridescence (Figs 24c-f); body punctate (Figs men was collected on Araucaria cunning- 24c-f); jugal margins of head carinate (Fig. hamii, but is treated here as a sitting record, 31b); metathoracic gland ostiole not hooded as this species is known to regularly cluster (Fig. 31d); peritreme narrow, elongate, arcu- on tall on-host trees during winter aestiva- ate, anteriorly-projected distally, medially sul- tion (Monteith, pers. comm.). cate (Fig. 31d); male pygophore caudally ori- Remarks: WHITE (1839) originally de- ented (Fig. 31e); spermathecal reservoir large, scribed C. parentum from an unspecified lo- oval; and, spermathecal bulb with proximal cation. SCHOUTEDEN (1906) described a sec- and distal flanges.

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Table 7: Choerocoris grossi nov.sp. and C. lattini nov.sp. diagnostic measurements in (Figs 24c-f), carinate in profile (Fig. 31b); millimetres. N = sample size. posterolateral margins rounded, carinate; Male Female posterior margin rectilinear. Scutellum: N Mean±SD Range N Mean±SD Range broadly U-shaped (Figs 24c-f), strongly con- Choerocoris grossi nov.sp. vex; forewing visible at base; connexiva vis- Length 5 10.60±0.64 10.15-11.05 5 10.72±0.61 9.70-11.22 ible. Thoracic Pleura: anterior margin of Pronotal width 5 5.45±0.61 5.03-5.88 5 5.76±0.40 5.07-6.05 proepisternum truncate to weakly explanate Width between eyes 5 2.33±0.23 2.17-2.50 5 2.47±0.16 2.21-2.60 (Fig. 31c); mesepimeron with evaporative Antennal segment length areas (Fig. 31d); external efferent system of I - 5 0.59±0.08 0.53-0.64 II - 5 0.44±0.12 0.36-0.53 metathoracic glands well-developed, cover- III - 5 0.56±0.09 0.52-0.59 ing most of metepisternum, raised (Fig. IV - 5 1.08±0.03 1.06-1.10 V - 5 1.15±0.09 1.08-1.21 31d); peritreme broad, sickle-shaped, anteri- Labial segment length orly directed distally, medially carinate (Fig. I 5 1.22±0.24 1.05-1.38 5 1.05±0.30 1.11-1.36 31d); evaporative areas barely surpassing II 5 1.33±0.10 1.26-1.40 5 1.33±0.15 1.11-1.50 peritreme distally, rugose (Fig. 31d). Pregen- III 5 1.28±0.31 1.06-1.50 5 1.21±0.17 0.94-1.33 IV 5 1.14±0.15 1.03-1.25 5 1.08±0.29 0.76-1.33 ital Abdomen: posterior angles of SIV-VII or III-VII weakly to strongly nodulate (Figs Choerocoris lattini nov.sp. 32a-h); males without sternal glands. Male Length 5 11.57±0.62 10.49-12.08 5 11.81±0.56 11.14-12.41 Genitalia: pygophore moderately sized, Pronotal width 5 5.99±0.21 5.77-6.24 5 6.13±0.22 5.90-6.38 lozenge-shaped, ventral margin rectilinear Width between eyes 5 2.74±0.16 2.50-2.89 5 2.74±0.09 2.67-2.85 to weakly excavate (Figs 31e, f, 32a, b, e-h, Antennal segment length 35a, 36a); genital opening narrowly oval, I 5 0.79±0.10 0.71-0.95 5 0.70±0.16 0.43-0.82 II(a) 5 0.44±0.06 0.35-0.50 2 0.43±0.33 0.38-0.48 dorsal in orientation, entire opening with II(b) 5 0.74±0.06 0.68-0.83 5 0.74±0.06 0.65-0.81 setose patches; parameres with short to III 5 0.63±0.20 0.46-0.92 5 1.01±0.15 0.82-1.16 elongate stem, small to moderately hooked IV 1 0.72 - 5 1.09±0.14 0.95-1.24 Labial segment length apex, sometimes with subdistal flange (Figs I 5 1.21±0.21 0.91-1.45 5 1.42±0.32 0.94-1.83 35b, 36b); ductus seminis proximalis nar- II 5 1.39±0.19 1.22-1.66 5 1.41±0.18 1.25-1.62 row, (Figs 35c, 36c); ejaculatory apparatus III 5 1.51±0.28 1.20-1.87 5 1.43±0.38 1.00-1.71 IV 5 1.37±0.10 1.28-1.51 5 1.40±0.31 1.22-1.52 moderately developed, with convoluted ventral conducting canal, 4-5 convolutions Description: Elongate-ovoid (Figs 24c- (Figs 35c, 36c); ejaculatory reservoir subel- f); strongly convex (Figs 32b, d, f, h); body liptoid, bag-like (Figs 35c, 36c); ductus sem- moderately sized, males 8-14.7 mm, females inis distalis and vesica greatly incrassate 10-14.5 mm; dorsum densely punctate (Figs subapically, vesica sometimes with subapi- 24c-f), sometimes less so on pronotum (Fig. cal, antler-like projections (Figs 36c, d). Fe- 24d); body glabrous (Figs 24c-f); appendages male Terminalia: small, in terminal plane; weakly setose; ground colour orange or red, paratergites VIII small and triangular; with fuscous to blue-fuscous markings, often paratergites IX large, subelliptoid; gonocox- with iridescence (Figs 24c-f); forewing not ae large, subtriangular, undivided, weakly surpassing tip of scutellum (Figs 24c-f). concave, medial margins weakly elevated. Head: moderately sized, suboval, transverse Spermatheca: short fecundation canal; (Fig. 31a); lateral margins of postclypeus sul- reservoir oval, internally ribbed; proximal cate; jugal margins sinuate, in profile cari- and distal flanges; bulb sclerotized, oval. nate (Fig. 31b). Antennae: AI-AII(b), AIV Diversity and distribution: Choerocoris is cylindrical, AIII flattened; AII(a) shortest an endemic Australian genus, which is com- segment, a little shorter than AI, AIV posed of four species, including two new longest segment. Labium: reaching between species, and are found commonly in conti- apices of metacoxae to abdominal sterna III; nental Australia (Fig. 34). LII longest segment, little longer than LIII & LIV (latter two subequal). Pronotum: Included species: broad, trapezoidal, strongly convex (Figs Ch. grossi nov.sp. Australia 24c-f); anterior margin weakly excavate; an- Ch. lattini nov.sp. Australia terolateral margins rectilinear to weakly Ch. paganus (FABRICIUS 1775) Australia convex, strongly to moderately divergent Ch. variegatus (DALLAS 1851) Australia

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Fig. 31: Scanning electron micrographs of key characters of Choerocoris paganus (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: C = clypeus; Ea = evaporative area; Pe = peritreme; Pe(k) = keel of anterior margin of proepisternum; P(r) = right paramere; Pr = proctiger. Scale bars = 1 mm.

Remarks: Choerocoris is a distinctive (1984) latter synonymized the genera, genus in the Scutellerinae, whose affinities adding that the female genitalia were also are uncertain. The ejaculatory apparatus is alike. On closer inspection, the aedeagus of typical of many scutellerines; with the ejacu- Calliscyta stalii shows close resemblance to latory apparatus possessing a convoluted Choerocoris paganus, in that the second con- ventral conducting canal and an enlarged junctival appendages are divided, with a conical lobal sclerite. However, this arrange- ejaculatory reservoir. It shares some similari- ment occurs in other scutellerines, and also ties with Cantao, in that the ductus seminis the elvisurine Solenotichus circuliferus. The distalis is incrassate subapically. It is similar two genera differ externally in that Choero- to many Australian scutellerines, with the coris has carinate jugal margins and a strong- first conjunctival appendages (CAI) absent. ly arcuate (and medially sulcate) peritreme MCDONALD (1961) first proposed that the of the metathoracic glands. male aedaegus of Calliscyta and Choerocoris were unlike, and MCDONALD & CASSIS

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Choerocoris grossi nov.sp. (Figs 24e, 32a, b, 33e, f, 34, Table 7) Holotype: Queensland: = Bunya Mountains, N Geary, 12 February 1940 (AM); Paratypes: 1Y, same data as holotype (AM); 1= 3YY, Rock- hampton, P1876-1877 (SAMA); New South Wales: 1= 1Y, Mullaley, 20 miles from Coon- abarabran, HJ Carter, November 1930 (AM; K63008); 1= 2YY, Mullaley, November 1957, FE Wilson (SAMA); 4YY 1 , Curlewis, A Musgrave, 29 October 1933, ex Beyeria viscosa (AM); 1Y, Mt Grattai, Range, H Paul, November 1933 (AM). Diagnosis: Choerocoris grossi nov.sp. is recognised by the following characters: yel- low, nodulate posterior angles of connexiva V-VII (Figs 32a, b); abdominal venter yel- low with lateral and submedial black mark- ings (Figs 32a, b); paratergites IX greatly en- larged, suboval (Fig. 33e), oblique in profile (Fig. 33f); gonocoxae I reduced, scythe-like, strongly incrassate medially (Figs 33e, f); and, male genitalia as in C. variegatus. Description: Body moderately sized, males 10.15-11.05 mm, females 9.70-11.22 mm. Colouration. Body bicoloured, dusty yel- low, with extensive black markings (Fig. 24c). Head: mostly black, clypeus with a lon- gitudinal yellow stripe (variable in length), lateral margins of jugae broadly yellow, rarely head mostly black, sometimes with green iri- descence; lorae black; genae mostly yellow, sometimes with dusty embrownment. An- tennae: black, sometimes with green irides- cence, rarely AI proximally orange-brown. Labium: mostly yellow, LIV fuscous. Prono- tum: mostly black, anterior and anterolater- al margins yellow, sometimes transverse yel- low stripe, submarginal to posterior margin (Fig. 24c). Scutellum: yellow anterior mar- gin with subhemispherical black marking, medially with W-shaped marking, sometimes contiguous with anterior marking, posterior margin with V-shaped black marking (Fig. 24c). Thoracic pleura and sterna: mostly yel- low, proepimeron and mesepimeron with black sublateral stripe, evaporative areas par- tially black. Legs: femora mostly black, with yellow markings, more so ventral surface; tibiae mostly black, sometimes with green iridescence, lateral surfaces yellow. Pregeni- tal Abdomen: posterior angles of connexiva Fig. 32: Colouration of Abdominal Venter of Choerocoris species (a-b) Choerocoris grossi V-VII yellow, nodulate (Figs 32c, d); abdom- nov.sp. (a) ventral view (b) lateral view (c-d) Choerocoris lattini nov.sp. (c) ventral view (d) lateral view (e-f) Choerocoris paganus (e) ventral view (f) lateral view (g-h) Choerocoris inal venter mostly yellow, lateral margins of variegatus (g) ventral view (h) lateral view. SIV-SVII with large, subtriangular black

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markings, submedial regions of SIV-SVII AII(a) smallest segment; AI & AII(b) sube- Fig. 33: Female Terminalia of Choerocoris with black suboval markings (Figs 32a, b); . qual in length. Labium: reaching posterior species (a) Choerocoris lattini nov.sp., ventral view (b) Choerocoris paganus, Male Genitalia: pygophore mostly yellow, margin of abdominal SIII; segments roughly ventral view (c-d) Choerocoris variegatus with margins black (Figs 32a, b). Female Ter- subequal in length, SII longest segment. Pre- (c) ventral view (d) lateral view (e-f) minalia: mostly black, paratergites IX with genital Abdomen: connexiva V-VIII with Choerocoris grossi nov.sp. (e) ventral view prominent yellow spots. posterior angles strongly incrassate, nodulate (f) lateral view. Abbreviations: PVIII = eighth paratergites; PIX = ninth (Figs 32a, b). Male Genitalia: ventral margin Texture. Dorsum densely punctate. paratergites; GXI = gonocoxae I. of pygophore deeply excavate; CAII(L) Structure. Antennae: females, AIV baglike with outer margins spiculate; heavi- longest segment, little longer than AIII; ly-sclerotized, antler-like CAIII; vesica with

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by the colouration of the abdominal venter, in possessing lateral and submedial bands of black colouration (cf. lateral band alone). The most conclusive diagnostic feature of C. grossi, is the female terminalia, which are unlike any species of Choerocoris, and in- deed any Australian species of jewel bug. In this species, paratergites IX are greatly en- larged, and much larger than gonocoxae I (cf. relatively small paratergites IX and large gonocoxae I), with the latter greatly re- duced, with the posterior margin deeply ex- cavate (Fig. 33e). Although the male geni- talia of C. grossi and C. variegatus are indis- tinguishable, the female terminalia serve as a strong basis for species separation.

Choerocoris lattini nov.sp. (Figs 24c, 32c, d, 33a, 34, 35a-d, Table 7) Holotype: Queensland: =, Bamaga, N Cape York, January 1958, PF Darlington, ex gum forest (AMNH). Paratypes: 1 larva, same data as holo- Y Fig. 34: Distribution of Choerocoris species. paired, subdistal, sclerotized processes. Fe- type (AMNH). 1 , Heathlands, 11°45’S 142°35’E, 1-6 April 1993, P Zborowski (ANIC); male Terminalia: paratergites VIII small, Northern Territory: 3== 3YY, Daly River, H subtriangular (Figs 33e, f); paratergites IX Wesselman (SAMA); 1Y, Pine Creek, Decem- greatly enlarged, suboval, oblique in profile ber 1908, B Hooper (SAMA); 1Y, Port Darwin (Figs 33e, f); gonocoxae I reduced, scythe- (SAMA); 1= 1Y, Cape York (SAMA); 1=, like, medially incrassate (Figs 33e, f). Cape York Penin[sula], C French, November 8 1892 (SAMA). Measurements. Table 7. Diagnosis: Choerocoris lattini nov.sp. is Etymology: This species is named after recognised by the following combination of Gordon Gross, in recognition of his major characters: head orange with subtriangular contribution to Australian Heteropterology. iridescent dark blue marking (Fig. 24c); ju- Distribution: This species is distributed gae and lorae orange; pronotum with orange in semi-arid regions of eastern Australia, transverse crescent-shaped fascia (Fig. 24c); from Rockhampton (Queensland) to loca- scutellum orange with broad anterior, paired tions west of the in subtriangular, and caudal V-shaped irides- New South Wales (Fig. 34). cent dark blue markings (Fig. 24c); AI or- ange; labium reaching abdominal sternum Host plants and biology: This species IV; abdominal venter mostly orange with has been collected on Beyeria viscosa. lateral subquadrate, blue markings (Figs 32c, Remarks: GROSS (1975) reported Choe- d); connexiva without yellow nodules (Figs rocoris variegatus to be composed of five sub- 32c, d); CAII(L) membraneous (Figs 35c, species; the nominotypical subspecies, C. d); CAII(M) mostly membraneous with api- variegatus similis, and three undescribed sub- cal sclerotization (Figs 35c, d); CAIII most- species. We have borrowed the material on ly membraneous, without pronounced which Gross made these conclusions, and processes (Figs 35c, d); vesica simple, mod- found a collection of specimens labeled as erately elongate, with subdistal tumescence ‘Choerocoris variegatus subsp.nov.’ These are (Fig. 35c); paratergites VIII small (Figs 32c, conspecific with a series of specimens of the d, 33a), triangular; paratergites IX small, Australian Museum, from New South Wales suboval (Figs 32c, d, 33a); and, gonocoxae I and Queensland. These specimens can be moderately sized, larger than paratergites IX distinguished externally from C. variegatus (Figs 32c, d, 33a).

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Description: Body moderately large, face sub-pentagonal (Fig. 35a); parameres males 10.49-12.08, females 11.14-12.41 with elongate stem, subdistal flange, and mm. short hook-shaped crown (Fig. 35b). phal- lotheca conical, with dorsal sclerotized areas Colouration. Body bicoloured, orange to (Figs 35c, d); ejaculatory apparatus with con- orange-red, with iridescent dark blue mark- voluted ventral conducting canal (Figs 35c, ings, sometimes with greenish tinge (Fig. d); elongate ejaculatory reservoir (Figs 35c, 24c). Head mostly orange-red with a subtri- d); ductus seminis distalis moderately incras- angular iridescent dark blue marking, widest sate (Figs 35c, d), without projections, ta- at posterior margin of head; lateral and ven- pered distally; vesica with out processes, tral aspects of head orange-red; eyes brown, moderately elongate, with subdistal tumes- ocelli red. Antennae: AI orange-red; AII(a)- cence (Fig. 35c); CAII(L) membraneous AIV dark brown. Labium: LI-III orange, (Figs 35c, d); CAII(M) mostly membraneous, stylets mostly fuscous; LIV dark brown to with apical sclerotization (Figs 35c, d); CAI- fuscous. Pronotum: anterior and anterolater- II mostly membraneous, lobe-like, medially al margins orange, remainder mostly irides- fused post-thecal margin (Figs 35c, d). Fe- cent blue, with a medial, transverse orange male Genitalia: paratergites VIII small (Figs crescent-shaped orange fascia, male with 32c, d, 33a), triangular; paratergites IX small, posterior margin a little orange (Fig. 24c). suboval (Figs 32c, d, 33a); gonocoxae I sub- Scutellum: mostly orange, with broad sub- triangular, weakly concave (Figs 32c, d, 33a). rectangulate iridescent dark blue marking on anterior margin, pair of subtriangular irides- Measurements. Table 7. cent dark blue markings, and caudal, small Etymology: This species is named after V-shaped, dark blue iridescent marking (Fig. Professor John D. Lattin of Oregon State 24c). Thoracic pleura and sterna mostly or- University, PhD supervisor of the senior au- ange, with iridescent dark blue or green thor [GC], in recognition of his knowledge markings on lateral margins of pleura; evap- of the jewel bugs of the world. orative areas grey. Legs: coxae and tro- chanters orange; femora mostly orange with Distribution: This species is known from small apical iridescent dark blue annulation; and the Northern Ter- tibiae and tarsi iridescent dark blue. Pregen- ritory (Fig. 34). ital Abdomen: venter mostly orange, with Host plants and biology: No host plant lateral margins of sterna with discontinuous, is known for this species. subquadrate, iridescent dark blue markings, SV-SVII with broad, V-shaped iridescent Remarks: Choerocoris lattini nov.sp. is dark blue marking; posterior angles of con- most closely related to C. variegatus, on the nexiva not swollen, fuscous (Figs 32c, d). basis of similarities of the male genitalia, particularly in the shape of the parameres Vestiture. Body mostly glabrous, with a and the conjunctival appendages (cf. Figs 35 few scattered setae on abdominal sterna, and 36), with the former species lacking more so on terminalia. antler-like CAIII. These species are howev- Texture. Dorsum uniformly punctate, er clearly differentiated by colour patterning with moderately deep, broad punctures (Fig. of the dorsum (cf. Figs 24c-f), the orange AI 24c); thoracic pleura and abdominal venter in C. lattini nov.sp. (dark brown in C. varie- densely punctate, with margins laterad of gatus), and the colour patterning of the ab- spiracular-trichobothrial region rugo-punc- dominal venter (cf. Figs 32c, d and g, h). tate. Choerocoris paganus (FABRICIUS 1775) Structure. Body oval, strongly convex. (Figs 1c, 24d, 31, 32e, f, 33b, 34) Antennae: AII(a) about 1/2 length of AI, Ground Jewel Bug shortest segment; AIII-AIV longest segment Cimex paganus FABRICIUS 1775: 698 (n.sp.); in females. Labium: reaching junction of ab- FABRICIUS 1781: 340 (description); FABRICIUS dominal sterna III/IV. Male Genitalia: dorsal 1787: 281 (description); FABRICIUS 1794: 84 (de- margin of pygophore arcuate, ventral margin scription); DONOVAN 1805: pl. 3 fig. 4 (descrip- emarginate (Fig. 35a); genital opening sur- tion)

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Fig. 35: Male genitalia of Choerocoris lattini nov.sp. (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII = third conjunctival appendage; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; F = flange at base of crown of paramere; GO = genital opening; S = stem of paramere; Sg = secondary gonopore; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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Tetyra pagana: FABRICIUS 1803: 134 (new combi- num orange to red. Pregenital Abdomen: nation); AMYOT & SERVILLE 1843: 47, pl. 1 fig. 6 venter mostly red to orange-red, with later- (description; habitus) al margins of SIV-SVII with triangular iri- Scutellera paganus: GUÉRIN 1831: pl. 11 fig. 5 (new descent green to blue-black markings (Figs combination); BOISDUVAL 1835: 625, pl. 11 fig. 4 32e, f); SIII-SVII with paired submedial (description); GUÉRIN-MÉNEVILLE 1838: 156 (de- oval dark markings (Figs 32e, f); SVII with scription) large, broad medial black marking (Figs 32e, Callidea pagana: GERMAR 1839: 122 (new combi- nation) f). Male Genitalia: pygophore uniformly Choerocoris paganus: DALLAS 1851: 29 (new com- black or red medially and remainder black. bination); VOLLENHOVEN 1863: 36 (description, Female Terminalia: most often red to or- distribution); MAYR 1866: 22 (description); STÅL ange-red, with minor black markings, to 1873: 13 (list); LETHIERRY &SEVERIN 1893: 21 paratergites VIII and IX and medial margins (catalogue); Distant 1899: 34 (biology); FROG- of gonocoxae I mostly black. GATT 1901: 1595 (description); FROGGATT 1902: 322 (description); SCHOUTEDEN 1904: 39 (list); Texture. Dorsum mostly densely punc- KIRKALDY 1909: 291 (catalogue); TILLYARD 1926: tate; pronotum with scattered punctures 149 (diagnosis); MCDONALD 1963a: 230, figs 1,2 (Fig. 24d). (female genitalia); MCDONALD 1963c: 290 (biol- Structure. Antennae: AII(a) shortest ogy); KUMAR 1964: 60 (male genitalia); GROSS 1975: 94, pl. C (description; colour habitus); segment, a little longer than AI; AIV MCDONALD &CASSIS 1984: 558, figs 61,62 (de- longest segment. Labium: reaching scription); CASSIS & GROSS 2002: 593 (cata- metasternum; LII longest segment, a little logue) longer than LIII and LIV. Male Genitalia: ventral margin of pygophore with broad, Diagnosis: Choerocoris paganus is recog- shallow notch; parameres with short hook- nised by the following combination of char- like crown, without sub-apical flange; phal- acters: scutellum mostly red, with pair of lotheca heavily sclerotized; ductus seminis submedial triangular blue markings (Fig. proximalis narrow, single membraneous 24d); callosite region of pronotum medially tube; ejaculatory apparatus with convoluted impunctate (Fig. 24d); callosite region pos- ventral conducting canal, 5-6 convolutions; teriorly demarcated by transverse furrow ejaculatory reservoir large, subelliptoid; (Fig. 24); abdominal venter orange with lat- base of ductus seminis distalis greatly in- eral and submedial dark blue markings (Fig. crassate; CAII mostly membraneous, bifid, 32e, f); CAII(M) and CAII(L) symmetrical, with small lobal sclerites; CAIII heavily with conical lobal sclerites; vesica with sub- distal tumescence; paratergites IX small, sclerotized, sinuate, distally acute, basally suboval (Fig. 33b); and gonocoxae I rela- fused (pre-thecal margin); vesica tapered tively large (Fig. 33b). distally without adornments. Female Termi- nalia: paratergites VIII moderate, size, sub- Description: Body moderately sized, triangular (Fig. 33b); paratergites IX small, males 8-11 mm, females 9-12 mm. suboval (Fig. 33b); gonoxocae I relatively Colouration. Body mostly red with iri- large, depressed (Fig. 33b); spermatheca as descent green-black to blue-black markings in generic description.

(Fig. 24d). Head: uniformly iridescent Measurements. MCDONALD &CASSIS green-black; underside of head iridescent 1984: Table 5. green to blue-black. Antennae: uniformly Y fuscous. Labium: fuscous. Pronotum: mostly Material examined: Queensland: 1 , Lizard Is- land, 29 September-1 October 1967, H Heatwole green to blue black, sometimes medially red (AM); New South Wales: 17== 9YY, Bundja- (Fig. 24d). Scutellum: mostly red, with sub- long National Park, Black Rocks, 32.24S medial pair of subtriangular green to blue- 152.32E, 15 November 1993, G Cassis, ex beach black iridescent markings, usually with an- wash (AM); 1=1Y, Mt Kaputar, Bullawa Creek, terior margin with large subrectangulate 28 April 1985, G Hangay (AM); 3== 1Y, green to blue-black iridescent marking, Goonoo State Forest, south side, 9 November sometimes absent and red (Fig. 24d). Tho- 1987, DK McAlpine & R de Keyzer (AM); Myall racic Pleura: nearly all iridescent green to Lakes National Park, 10.3 km S Seal Rocks Road blue-black, with anterior edge of proepister- on Hawks Nest Road, 32°30’S 152°21’E, October

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Fig. 36: Male genitalia of Choerocoris variegatus (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII = third conjunctival appendage; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; F = flange at base of crown of paramere; GO = genital opening; S = stem of paramere; Sg = secondary gonopore; V = vesica; Vp = process of vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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20 1995, RT Schuh & G Cassis, ex Dodonaea vis- tered jewel bugs in Australia. It can be read- cosa angustissima (AM); 3==, 2.9 km W Nyn- ily identified on colour patterning alone; gan, 31°33’56“S 147°09’19“E, 202 m, October with the dorsum mostly red with iridescent 18.2001. RT Schuh, G Cassis, R Silveira & MA blue-black markings. It can also be separat- = Wall, ex Dodonaea viscosa spatulata (AM); 1 , ed from its congeners by texture, with the Munmorah State Recreation Area, 33°12.26’S pronotum less densely punctate; the cal- 131°34.37’E, 11 October 1997, L Wilkie, losite region is medially impunctate (cf. uni- K145365 (AM); 1Y, Booti Booti National Park, formly and densely punctate in others). The 32°11.15’S 152°31.42’E, 25 November 1997, L Wilkie (AM); 5== 9YY, Yara, 32°51’54“S male genitalia of C. paganus are also dis- 146°11’21“E, December 2000, G Swan, semiarid tinct, with the CAII bifid, each branch pos- grassland, pitfall trap (AM); 2== 4YY, Yara, sessing a conical lobal sclerite. 32°56’48“S 146°11’32“E, December 2000, G Swan, spinifex grassland, pitfall trap (AM); 4== Choerocoris variegatus DALLAS 1851 11YY, Yara, 32°56’39“S 146°11’32“E, Novem- (Figs 24f, 32g, h, 33c, d, 34, 35) ber-December 1997, G Swan & E Wapstra, Choerocoris variegatus DALLAS 1851: 29 (n.sp.); mallee woodland, pitfall trap (AM); Southern STÅL 1873: 13 (list); LETHIERRY & SEVERIN 1893: Australia: 1=, 7 km E of Para Wirra National 21 (catalogue); SCHOUTEDEN 1904: 39 (list); Park, nr. Williamstown, 34.42S 138.51E, 31 Oc- KIRKALDY 1909: 292 (catalogue); TILLYARD 1926: tober 1995, RT Schuh, G Cassis, & GF Gross, 149 (note); MCDONALD 1963a: 30 (male geni- Site 95-39. (AM); Western Australia: 9== talia); MCDONALD 1963b: 230, figs 3-5 (female 7YY, Frank Hann National Park, Lillian Stoke genitalia); MCDONALD 1963c 285 (larvae); KU- Rock, 33.066S, 120.083E, 400 m, 5 November MAR 1964: 60-61, 63 (morphology); KUMAR 1996, G Cassis & RT Schuh, Site 96-66, ex 1965: 44, 52-53 (male genitalia); MCDONALD & Dodonaea viscosa angustissima (AM); Northern CASSIS 1984: 559 (description); MCDONALD Territory: 4== 3YY, 13.5 km E Stuart High- CASSIS & GROSS 2002: 594 (catalogue) way, on Horseshoe Bend Road, 464 m, Choerocoris similis DISTANT 1899: 34 (n.sp.); 28°08’53“S 133°17’59“E, 28 October 2001, G FROGGATT 1907: 327 (note); MCDONALD & CAS- Cassis, RT Schuh, MD Schwartz, R Silveira & SIS 1984: 559 (synonymy) MA Wall, ex Dodonaea viscosa (AM); Trephina Choerocoris variegatus similis: GROSS 1975: 95 Gorge National Park, John Hayes Rockpool (subspecies) Campground, 580 m, 23°23’30“S 134°21’15“E, Diagnosis: Choerocoris variegatus is 25 October 2001, G Cassis, RT Schuh, MD recognised by the following characters: Schwartz & R Silveira & MA Wall, ex Dodonaea pronotum uniformly punctate (Fig. 24f); viscosa mucronata, Site CA01L21H79 (AM). body yellow to red, with black markings Distribution: This species is broadly dis- (Fig. 24d); posterior angles of connexiva V- tributed in Australia, and is known from all VIII not expanded (Figs 32g, h); abdominal states and territories, aside from the Aus- venter yellow with lateral black markings tralian Capital Territory (Fig. 34). (submedial markings lacking) (Figs 32g, h); stem of parameres elongate (Fig. 36b); Host plants and biology: This species is CAII(L) with basal denticulation (Figs 36c, known primarily from a number of sub- d); CAIII large, sclerotized, antler-like (Figs species of Dodonaea viscosa (Table 1). It has 36c, d); vesica with pair of subapical also been collected on a variety of plants, processes (Fig. 36c); and, gonocoxae I mod- and appears to be polyphagous, although erately developed, larger than paratergites these plants are likely to be secondary food- IX (Figs 33c, d). preferences. MCDONALD (1960, 1963c) de- scribed its biology, and found that they are Description: Body moderate size, males both pre- and post-dispersal seed predators. 8-12 mm, females 10.2-12.9 mm. He also reported it often found amongst Colouration. Body bicoloured, dusty rocks, within the ‘seed shadow’ of their host yellow to red, with extensive black mark- plants. The adults are known to overwinter ings, not iridescent (Fig. 24f). Head: mostly in dry places, and the larvae are sometimes black, clypeus with a longitudinal yellow found in grass tussocks. stripe (variable in length), lateral margins of Remarks: Choerocoris paganus is one of jugae yellow, sometimes jugae with small the most distinctive and commonly encoun- medial yellow spot to yellow longitudinal

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stripe; lorae black (sometimes with yellow tized, large, antler-like (Figs 36c, d). Female spot); jugae yellow. Antennae: mostly black, Terminalia: paratergites VIII subtriangular with AI partially to mostly yellow; remain- (Figs 33c, d); paratergites IX suboval, mod- der fuscous. Labium: mostly yellow, LIV fus- erately sized, weakly incrassate medially cous. Pronotum: mostly black, anterior, an- (Figs 33c, d); gonocoxae I large, depressed terolateral and posterior margins mostly yel- medially, medial margins recurved (Figs 33c, low (punctures black), with medial pair of d); spermatheca as in generic description. yellow or red elliptoid markings, sometimes Measurements. MCDONALD &CASSIS almost contiguous medially (Fig. 24f). 1984: Table 5. Scutellum: yellow to red, anterior margin with subrectangulate black margin, medially Type material examined: Choerocoris variegatus with W-shaped marking, sometimes con- DALLAS: Holotype, Y, ‘Swan River’, ‘B.M. Hem. tiguous with anterior marking, posterior Type No. 426’ (BMNH); Choerocoris similis DIS- Y margin with V-shaped black marking (Fig. TANT: Holotype, , ‘Adelaide’ ‘B.M. Hem. Type 24f). Thoracic Pleura: mostly yellow, No. 427’ (BMNH). There are two other speci- mens in the BMNH of Choerocoris variegatus, proepimeron with black sublateral stripe, which are labeled as paratypes, and have the evaporative areas partially black. Legs: same locality as the holotype. DALLAS (1851) on- forefemora bicoloured, yellow, with distal ly referred to a single specimens (as ‘a. Swan Riv- regions blackened (sometimes with black er’), and we interpret this is the holotype, and spotting), dorsal surface of meso- and have no evidence for designating a lectotype. metafemora black, remainder mostly yellow, Other material examined: New South Wales: tibiae mostly black, laterally with yellow 1=, Round Hill Fauna Reserve, 9 April 1977, G stripes; tarsi black. Pregenital Abdomen: Daniels (AM); 1=, Round Hill Reserve, near Eu- venter mostly yellow, lateral margins of SIV- abalong, 28 December 1992, MS & BJ Moulds SVII with large, subtriangular black mark- (AM); 1=, 30 km E Southern Cross, 30 Septem- ings (Figs 32g, h). Male Genitalia: py- ber 1985, J Bugeja (AM); Western Australia: 1Y, 115.4 km E of Norseman, 32.05S 122.966E, gophore mostly yellow, with margins black. 600 m, RT Schuh & G Cassis, 23 October 1996, Female Terminalia: mostly black, with me- Site 96-10, ex Beyeria lechenaultii; 9== 2YY, dial regions of gonocoxae I and paratergites Duke of Orleans Bay, Table Island Picnic Area, with yellow markings. 33.899S 122.594E, 50 m, 24 November 1999, RT Structure. Antennae: AI and AII(b) Schuh, G Cassis & R Silveira, Site 99-32, ex Spyridium globulosum (AM); 2== 7YY, Duke of subequal in length; AII(a) shortest segment; Orleans Bay, E Esperance, 14 November 1993, J AIII and AIV longest segment. Labium: & A Leask (AM); 1= 2YY, 5 km W Wuarga, 2 reaching apex of metacoxae; LI shortest seg- September 1981, GA Holloway (AM); 5==, ment; LII-LIV subequal in length, some- Rossiter Bay, Cape Le Grande National Park, times LII a little longer than LIII and LIV. 33°58.0345’S 122°16.0457’E, 3 m, 23 November Male Genitalia: pygophore lozenge-shaped, 1999, RT Schuh, G Cassis & R Silveira, Site 19- with narrow genital opening, ventral margin 29, ex Acacia cyclops (AM). deeply excavate (Fig. 36a); parameres with Distribution: This species is broadly dis- stem greatly elongate, with lateral flange, tributed in Australia, and is known from New and weakly hooked crown (Fig. 36b); phal- South Wales, Northern Territory, Queens- lotheca conical, with ventral areas of sclero- land, South Australia, Victoria, and Western tisation (Fig. 36c); ductus seminis proxi- Australia. It is primarily known from temper- malis narrow; ejaculatory apparatus well de- ate Australia, although a few specimens have veloped, with convoluted ventral conduct- been taken in tropical Queensland and the ing canal, 6-8 convolutions (Fig. 36c); ejac- Northern Territory (Fig. 34). ulatory reservoir suboval, moderately sclero- tised (Fig. 36c); CAII mostly membraneous, Host plants and biology: The biology of bifid, CAII(L) with basal denticulation, Choerocoris variegatus is not well-known. It CAII(M) with conical lobal sclerite (Figs has been collected on numerous occasions 36c, d); ductus seminis distalis not greatly in association with seeds, mostly on plants. incrassate (Fig. 36c); vesica with arcuate It has been collected on a number of plants subapical processes, secondary gonopore (Table 1), but its primary hosts are thought apical (Figs 36c, d); CAIII heavily sclero- to be species of the euphorb genus Beyeria.

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Remarks: Choerocoris variegatus is wide- eas extensive, rugose, extending beyond per- spread, but is less commonly encountered itreme and on to mesepimeron (Fig. 38c); os- than its congener, C. paganus. It has male tiole small; peritreme raised, obovate (Fig. genitalia varying only in shape and size of 38c). Abdominal Venter: SIII carinate along the CAIII; the aedeagus cannot be separat- midline. Female Terminalia: paratergites IX ed from those of the new species, C. grossi small, subelliptoid; gonocoxae I large, subtri- nov.sp. Differences in colour patterns and angular, posterior margin weakly excavate. female terminalia are sufficient to separate Spermatheca: basal sclerites present; fecun- species. dation canal very short (Fig. 38d); reservoir oval (Fig. 38d); prominent pump (Fig. 38d). Heissiphara nov.gen. (Figs 24b, 27, 37, 38) Male unknown. Type species: Heissiphara minuta nov.sp., original Etymology: This genus is named after designation our colleague Dr. Ernst Heiss in recognition of his significant contribution to Hetero- Diagnosis: Heissiphara nov.gen. is recog- pterology. nised by the following combination of char- acters: small species (Figs 24b, 37); dorsum Included species: strongly convex; dorsum heavily punctate H. minuta nov.sp. Australia with setigerous punctures (Figs 24b, 37); head strongly declivent and rounded (Fig. Diversity and distribution: Heissiphara is 38b); jugae rounded (Fig. 38b); bucculae a monotypic genus, known from a single lo- small, arcuate (Fig. 31c); anterior margin of cality in temperate Western Australia. scutellum with submedial subreniform pol- Remarks: Heissiphara superficially re- ished calli (Fig. 24b, 37); lateral margins of sembles a number of Australian pentatomid pronotum subcarinate; external efferent sys- species, with an enlarged scutellum, such as tem of metathoracic glands moderately de- Kapunda. However, Heissiphara is a member veloped, with raised obovate peritreme (Fig. of the Scutelleridae based on the following 38c); evaporative areas extending to characters: greatly enlarged scutellum, sper- mesepimeron (Fig. 38c); spermathecal fe- matheca with basal sclerites, and the sper- cundation canal very short (Fig. 38d); reser- mathecal reservoir without a sclerotized rod. voir oval, not heavily sclerotized; and, sper- mathecal pump large (Fig. 38d). The position of Heissiphara within the scutellerid hierarchy is difficult to determine, Description: Small species (Figs 24b, as males are unknown. It is conclusively ex- 37); ovoid; dorsum strongly convex (Figs cluded from the Elvisurinae in that it does 24b, 37); body evenly punctate, with setiger- not possess keel-like thoracic sterna. It is un- ous punctures (Figs 24b, 37). Head: trans- likely to be a member of the Tectocorinae as verse (Figs 38a), strongly rounded and de- its female terminalia and spermatheca are clivent (Fig. 38b); lateral margins of jugae consistent with those of the scutellerines. rounded (Fig. 38b). Antennae: AIII and Heissiphara shares some similarites with AIV bicompressed. Pronotum: subtrape- Morbora, in that both the body has setigerous zoidal, strongly convex (Figs 24b, 37); an- punctures and the metathoracic peritreme is terolateral margins elongate, rectilinear, sub- obovate. However, Heissiphara also shares carinate; posterolateral margins short, these characters with Choerocoris, and on the rounded; posterior margin truncate (Figs basis of the similar spermathecae, the former 24b, 37). Scutellum: broad, U-shaped, base genus is putatively assigned to the Scutelleri- of forewings and abdominal connexiva III- nae, despite its dull colouration. VII visible (Figs 24b, 37); anterior margin with submedial, subreniform, polished calli Heissiphara differs from Choerocoris in (Figs 24b, 37); tip of forewings not exposed; the shape of the head, which is more round- propleuron large, platelike, with anterior ed and the lateral jugal margins are not ex- margins of proepisternum arcuate; external planate. In addition, Heissiphara has a more efferent system of metathoracic glands mod- densely punctate body, and is significantly erately developed (Fig. 38c), evaporative ar- smaller, the anterior margin of the scutellum

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yellow with brown apex and fuscous mid- line; eyes reddish brown; ocelli yellow. An- tennae: AI-AII(a&b) yellow, AIII and AIV dark brown. Labium: LI and II yellow, LIII and LIV brown. Pronotum mostly light yel- low-brown to medium brown, with darker punctations; callosite region anteriorly with yellow fascia and calli yellow; disc with me- dial rectangular red mark and two adjacent submedial fuscous spots. Scutellum: ground colour mostly concolorous with pronotum, anterior margin with central red marking and two yellow submedial calli; anterolater- al areas with dark brown foveae; posterolat- erally with dense patches of dark puncta- tions forming obscure semi-circular mark- ing. Thoracic pleura and abdominal venter uniformly yellow with brown punctations. Legs uniformly yellow-brown with darker brown punctations. Vestiture. Body glabrous. Antennae: AI-AII(a&b) with sparse distribution of short setae; AIII-AIV with denser distribu- tion of setae. Texture. Body densely punctate, with setiferous punctures. Structure. Head: interocellar space twice distance between ocelli and eye. An- tennae: AII(a) and AII(b) short, subequal Fig. 37: Habitus of holotype of Heissiphara has submedial calli, and the spermathecal in length; AIV longest segment. Labium: Y minuta nov.sp., . reservoir is not heavily sclerotised. reaching metasternum; LI and LII cylindri- cal; LIII and IV flattened. Abdominal seg- Heissiphara minuta nov.sp. ment III with a medial groove to accommo- (Figs 24b, 27, 37, 38) date rostrum; apical margins of segments III Holotype: Y, Western Australia: Coastal High- to VI tumescent. Female genitalia: see way, 57 km N of Kalbarri Road, -27.26.853S, generic description. 114.41.204E, altitude 500m, G. Cassis and R.T. Male unknown. Schuh ex Stenanthemum complicatum (AM); Paratype: Y, same data as holotype (AM). Measurements. Holotype: BL: 3.35, PW: 2.39, IOD: 1.12, AI: 0.26, AII(a): 0.16, Diagnosis: This species is recognised by AII(b): 0.18, AIII: 0.30, AIV: 0.40, LI: 0.49, the following combination of characters: LII:0.63, LIII: 0.39, (LIV not observable). small size, < 3.5 mm; body strongly convex (Fig. 24d); body uniformly distributed with Etymology: This species is named for its setigerous punctures (Fig. 24d); AII(b) small size. shortest antennal segment; pronotum and Distribution: Heissiphara minuta nov.sp. scutellum with medial red markings (Fig. is known from the type locality only, north 24d); and, female terminalia and spermath- of Kalbarri National Park, on the central- eca as in generic description. west coast of Western Australia (Fig. 27). Description: Colouration. Head: mostly Host plants and biology: This species yellow; posterior margin of vertex with tri- was collected from Stenanthemum complica- angular fuscous markings; clypeus mostly tum (Rhamnaceae).

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Fig. 38: Scanning electron micrographs of key characters of Heissiphara minuta nov.sp. (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) Spermatheca. Abbreviations: Df = dorsal flange; Ea = evaporative area; Pe = peritreme; Pf = proximal flange; P(r) = right paramere; S(b) = spermathecal bulb. Scale bars = 1 mm.

Remarks: Heissiphara minuta nov.sp. is Diagnosis: Lampromicra is recognised by one of the smallest species of scutellerid the following combination of characters: known to us, rivalling odontotarsine species moderately sized-body; iridescent coloura- like Odontoscelis signatus FIEBER, O. tomentosa tion Figs 1d, 28c-e); dorsum and abdominal (GERMAR) and Psacasta lethierryi PUTON. Un- venter setose (Figs 1d, 28c-e); lateral mar- like most other scutellerines, H minuta gins of jugae weakly sinuate (Fig. 39a); an- nov.sp. is relatively dull in colouration; most- tennae short; AII(a) a little shorter than AI; ly pale brown, with yellow markings. It is AI-AII(a&b) rounded; AIII-AIV flattened; known only from females. AIV longest segment; scutellum with tumescent anterior callus, impunctate, Lampromicra STÅL 1873 (Figs 1d, 21f- rarely punctate; forewing extending beyond h, 28c-e, 39, 40, 41, Table 8) scutellum (Figs 28c-e); proepisternum weak- Philia SCHIÖDTE 1842: 279 (gen. nov.) (junior ly explanate (Fig. 39c); external efferent sys- homonym of Philia MEIGEN 1800 [Diptera]); STÅL tem with a large ostiole; broad, subreniform 1868: 9 (description); LETHIERRY & SEVERIN peritreme, without a medial, longitudinal 1893: 22 (catalogue); SCHOUTEDEN 1904: 28 (de- groove (Fig. 39d); evaporative areas extend- scription); ing to mesepimeron (Fig. 39d); ventral bor- Philya STÅL 1865: 33 (incorrect subsequent der of pygophore rounded (Fig. 39e) to spelling; key); STÅL 1866: 151 (incorrect subse- emarginate; dorsal margin of male genital quent spelling) opening with setal patches (Fig. 40a); para- Lampromicra STÅL 1873: 16 (gen. nov.); meres with moderately sized hook-shaped KIRKALDY 1909: 299 (synonymy); MCDONALD & apex (Fig 40b); phallotheca with pair of sub- CASSIS 1984: 555 (description); CASSIS & GROSS 2002: 595 (catalogue) distal, thornlike processes (Fig. 40c); ductus Schioedtia KIRKALDY 1905: 79 (nom. nov. for Phil- seminis distalis S-shaped (Figs 40c, d); ven- ia SCHIÖDTE 1842) tral conducting canals of ejaculatory appara- Type species: Tetyra senator FABRICIUS 1803, sub- tus, with ≤ 10 convolutions (Figs 40c, d); sequent designation, SCHOUTEDEN 1904: 29 CAI absent; CAII bifurcate, CAII(M)

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Table 8: Lampromicra aerea: diagnostic measurements in millimetres. N = 8 males and 8 weakly convex; posterior margin rectilinear females. to weakly convex (Figs 28c-e). Scutellum: Male Female V-shaped, strongly tapered posteriorly, Mean±SD Range Mean±SD Range strongly declivent beyond connexiva V, pos- Length 11.58±0.52 11.00-12.17 11.46±0.34 11.00-11.50 terior margin subtruncate (Figs 28c-e). Tho- Pronotal Width 6.33±0.76 5.12-7.08 6.15±0.54 5.67-6.92 racic pleura: anterior margin of proepister- Width between the eyes 2.21±0.05 2.17-2.28 2.22±0.07 2.08-2.32 num weakly explanate (Fig. 39c); Antennal segment length I 0.63±0.14 0.60-0.68 0.87±0.04 0.80-0.92 mesepimeron with evaporative areas (Fig. II(a) 0.64±0.04 0.60-0.68 0.65±0.03 0.60-0.68 39d); external efferent system of metatho- II(b) 1.32±0.07 1.24-1.40 1.28±0.05 1.20-1.32 racic glands well-developed (Fig. 39d); osti- III 1.80±0.06 1.72-1.84 1.69±0.21 1.60-1.72 IV 1.96±0.04 1.92-2.00 1.90±0.08 1.80-2.00 ole large, hooded; peritreme raised, broadly Labial segment length subreniform (Fig. 39d); evaporative areas of I 0.96±0.03 0.92-1.16 0.93±0.04 0.88-1.16 metepisternum extending little beyond per- II 1.83±0.22 1.60-2.04 1.63±0.07 1.56-1.84 III 1.34±0.04 1.24-1.40 1.29±0.04 1.24-1.32 itreme (Fig. 39d). Pregenital Abdomen: pos- IV 1.34±0.80 1.32-1.40 1.28±0.05 1.20-1.32 terolateral angles without tubercles (Figs 21f-h). Male Genitalia: pygophore with basally membraneous, with hook-shaped lobal sclerite (Fig. 40c); CAII(L) membra- ventral surface concave, ventral margin neous, with denticulate apex, perpendicular rounded (Figs 39e, f, 40a) to excavate; gen- to hook-shaped (Figs 40c, d); vesica S- ital opening with dorsal setal patches (Fig. shaped (Fig. 40c, d); spermathecal fecunda- 40a); parameres with columnar base, and tion canal elongate; and, spermathecal relatively short, hook-shaped apex (Fig. reservoir broad, oval. 40b); phallotheca box-like, with pair of small, subdistal, thorn-like processes (Fig. Description: Body elongate-ovoid, 40c); ejaculatory apparatus with elongate strongly tapered caudally (Figs 28c-e); mod- ventral conducting canals, ≤ 10 convolu- erately-sized body, males 9-12 mm, females tions (Fig. 40c, d); ejaculatory chamber 8.8-13 mm; iridescent colouration, most of- elongate (Fig. 40c); dorsal conducting canal ten dark blue or green (Figs 1d, 28c-e), broad; ductus seminis and vesica distalis S- sometimes orange, often with contrasting shaped (Fig. 40c); CAII bifurcate, CAII(M) markings on pronotum and scutellum, ei- basally membraneous, with hook-shaped ther black, orange, bronze or green; body lobal sclerite (Fig. 40c); CAII(L) membra- strongly setose (Figs 1d, 28c-e); dorsum with neous, with denticulate apex, perpendicular dense distribution of shallow to moderately to hook-shaped (Figs 40c, d). Female Termi- deep punctures (Figs 28c-e), sometimes nalia: paratergites VIII moderately-sized, more scattered on pronotum; ventral surface subtriangular; paratergites IX moderately- of body and appendages densely setate, less sized, subelliptoid; gonocoxae I large, subtri- so on thoracic pleura. Head: subtriangular angular, outer surface concave. Spermathe- (Fig. 39a), transverse, weakly declivent; lat- ca: fecundation canal elongate; spermathe- eral margins of jugae weakly excavate (Fig. cal reservoir broad, oval; pump well-devel- 39a), subcarinate in profile (Fig. 39b). An- oped, with proximal and distal flanges. tennae: AII(a) a little shorter than AI; AI- Diversity and distribution: Lampromicra AI(a&b) rounded; AIIII-AIV flattened; comprises 17 species, three of which occur AIV longest segment. Labium: reaching be- in Australia (aerea, regia and senator). It also tween the apices of the metacoxae to the occurs in Indonesia (6 species), New mid-point of abdominal sternite IV; LII Guinea (7), Philippines (2), New Caledonia longest segment; LIII and LIV roughly sube- (4), and the Solomon Islands (2), with a qual in length. Pronotum: moderately con- number of these species occurs in at least vex; anterior margin weakly excavate; an- two of these countries. Two of the Aus- terolateral margins rectilinear, strongly di- tralian species have restricted distributions, vergent, carinate in profile (Fig. 39b); cal- whereas L. senator is in tropical north Aus- losite region demarcated posteriorly by tralia, extending on the eastern coast as far transverse trough-like depression, some- south as the Northern Rivers of New South times punctate; posterolateral margins Wales (Fig. 41).

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Included species: Lampromicra aerea DISTANT 1892 L. aerea (DISTANT 1892) Australia (Figs 21f, 28c, 40, 41, Table 8) Revised L. balteata (WALKER 1867) New Guinea Status

L. cuprina (STÅL 1873) New Guinea Philia aerea DISTANT 1892: 96 (n.sp.); SCHOUTE- L. caledonica (DISTANT) New Caledonia DEN 1904: 29 (list) L. distinguenda (WALKER 1868) Indonesia Philia compacta BREDDIN 1903: 57 (n.sp.); L. ditissima (VOLLENHOVEN 1863) SCHOUTEDEN 1904: 29 (synonymy) Indonesia, New Guinea Lampromicra aerea: KIRKALDY 1909: 299 (new L. elegans (MONTROUZIER 1861) combination); MCDONALD 1963: 26 (male geni- New Caledonia talia); MCDONALD & CASSIS 1984: 556 (as junior L. fastuosa (VOLLENHOVEN 1863) synonym of Lampromicra senator); CASSIS & Indonesia, New Guinea GROSS 2002: 597 (catalogue; as junior synonym of Lampromicra senator) L. festiva (GERMAR 1839) Philippines L. geminata (DISTANT) New Caledonia Diagnosis: Lampromicra aerea is recog- L. geniculata (STÅL 1871) Philippines nised by the following combination of char- L. jactator (STÅL 1854) Indonesia, acters: dorsum iridescent copper-fuscous New Guinea (Fig. 28c); femora red; abdominal venter L. leucocyanea (MONTROUZIER 1855) fuscous, with lateral regions red (inner mar- New Caledonia, New Guinea, gin of stripe linear) (Fig. 21f); dorsum setose Solomon Islands (Fig. 28c); anterior callosite region of L. regia (BERGROTH 1895) Australia scutellum punctate (Fig. 28c); and, CAII L. senator (FABRICIUS 1803) Australia, dorsal branch with hooked, denticulate lob- New Guinea, Indonesia al sclerite (Figs 40c, d). L. vulcanica (LE GUILLOU 1841) Indonesia L. woodfordi (DISTANT 1899) Solomon Description: Moderate size species, males Islands 11.0-12.17 mm, females 11.0-11.5 mm.

Remarks: There has been no modern Colouration. Dorsum uniformly copper- comprehensive systematic treatment of fuscous, often with purple iridescence (Fig. Lampromicra. MCDONALD & CASSIS (1984) 28c); eyes and ocelli red; antennae fuscous; redescribed the genus on the basis of the labium mostly fuscous, with LI burnt-orange Australia species, highlighting the impor- highlighting. Thoracic pleura and sterna tance of the male genitalia. LYAL (1979) uniformly black. Legs: coxae and trochan- mentioned the diagnostic proximity of Lam- ters burnt-orange with dusty black high- promicra with Calliphara and Chrysophara. lighting; femora red; tibiae mostly fuscous Lampromicra differs from Calliphara in hav- with narrow basal red annulation; tarsi ing a strongly setose dorsum (Fig. 1d), black. Pregenital Abdomen: venter mostly trough-like posterior margin of the callosite fuscous, with region lateral to spiracular-tri- region of the pronotum (Figs 28c-e), round- chobothrial region red, sometimes orange- ed connexival angles (Figs 21f-h), the CAI red, inner margin of stripe linear (Fig. 21f). absent (Figs 40a, d), CAII(L) perpendicular Vestiture. Body with dense distribution to hook-shaped with denticulations (Figs of elongate, fine, erect setae (Fig. 28c). 40a, d), and the vesica is S-shaped (Figs 40a, d). The setose dorsum is not useful for ge- Texture. Body with dense distribution neric boundaries, as it occurs in other scu- of shallow punctures. Pronotum: indistinct tellerine taxa, such as species of Brachaulax, rows of deeper punctures on anterior and Procilia, Scutellera and Tetratharia, as well as posterior margins of callosite region (Fig. the Australian species, Cantao parentum. 28c). Scutellum: anterior callosite region punctate; lateral margins weakly rugopunc- The species taxonomy of Lampromicra is tate. Abdominal Venter: region lateral of exceedingly complex, with high intra-popu- spiracular-trichobothrial region rugopunc- lation variation in body colour and shape of tate (Fig. 21f). the male conjunctival appendages. The lob- al sclerite of CAII(L) exhibits continuous Structure. Antennae: AI and AII(a) variation; particularly in the ubiquitous subequal in length, AI little longer in fe- species, L. senator. males; AIV longest segment. Labium: ex-

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Fig. 39: Scanning electron micrographs of key characters of Lampromicra senator (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Scale bars = 1 mm.

tending to junction of abdominal sterna appendages in resting position (Figs 40c, d); III/IV. Pronotal width broader than 1/2 body CAI absent (Figs 40c, d); CAII(M) with length. Head: weakly convex; broad. Male conical lobal sclerite (Figs 40c, d); CAII(L) Genitalia: pygophore with broad setal patch membraneous, with large hook-shaped, den- on dorsal margin of genital opening (Fig. ticulate lobal sclerite (Figs 40c, d); CAIII 40a); parameres with short hook-shaped arcuate, heavily sclerotised, separated post- thecal margin (Fig. 40d). Female Genitalia: apex (Fig. 40b); phallotheca with moderate- paratergites VIII small, subtriangular; ly large subapical thornlike processes (Fig. paratergites IX small, subelliptical; outer 40b); ejaculatory apparatus with prominent surface of gonocoxae I weakly concave. ventral conducting canals, up to 10 convo- Spermatheca: fecundation canal elongate; lutions (Figs 40c, d); ejaculatory reservoir reservoir, large, oval; pump well-developed. elongate (Fig. 40c); dorsal conducting canal broad; ductus seminis distalis and vesica S- Measurements. See Table 1. shaped, extending to apex of conjunctival

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Fig. 40: Male genitalia of Lampromicra aerea (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII = third conjunctival appendage; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; Pt(P) = process of phallotheca; S = stem of paramere; Sg = secondary gonopore; SP = setose patch; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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Type material examined: Philia aerea DISTANT: or 1/3 of scutellum purple-fuscous, with iri- Holotype, Y, ‘Sidney’, ‘B.M. Hem. Type No. descent green tinge (Fig. 28d); posterior 2/3 456’, ‘Dist. Coll. 1911-383’ (BMNH). of scutellum yellow (Fig. 28d); femora yel- Other material examined: New South Wales: low; tibiae iridescent green; abdominal ven- 1=, Woody Head, near Iluka, 31 December 1978, ter mostly dark iridescent green, lateral mar- BJ Day, ex rainforest (AM); 1Y, 3 km NE Har- rington, 30 November 1987, G Williams, ex lit- gins green (Fig. 21g); CAII(M) apex serrate toral rainforest (AM); 1Y, Coory, P Cantwell, 7 and hooked; CAIII medially fused; and, January 1983 (AM); 1=, Mooney Mooney Creek, vesica moderately sized, weakly arcuate. near Gosford, 10-11 November 1982, B Day & K Description: Body moderate-sized, males Khoo (AM); 1Y, 17 km N Macksville, 4-xii- 9.3-12.0 mm, females 8.8-12.0 mm. 1948 (BMNH); 1Y, Cabramatta, George’s River Valley, 5-30-I-1963, M Nikitin, B.M. 1963-283, Colouration. Body bicoloured (Fig. = on flowers of (BMNH); 1 , Syd- 28d), with head, pronotum and anterior 1/2 ney, x-1904, AP Dodd, ‘H65’, B.M. 1923-124 of scutellum (latter (pentagonal marking) (BMNH); 1Y, Nowra, FA Rockway, B.M. 1929- purple-fuscous with green iridescence, re- 45 (BMNH); 1Y, Hornsby, 19-xi-1969, MI Nikitin (BMNH); 1Y, Lane Cove, 23-x-1958, mainder of scutellum yellow; coxae, B.M. 1964-57 (BMNH); 1=, Cabramatta, trochanters and femora yellow, tibiae most- George’s River Valley, MI Nikitin, B.M. 1962- ly iridescent green with bases narrowly yel- 347, ex Kunzea ambigua (BMNH). low; tarsi iridescent green; abdominal venter Distribution: Lampromicra aerea is known mostly iridescent green, lateral margins yel- primarily from the Basin, with its low (Fig. 21g). most northern limit near Iluka on the North- Texture. Head sparsely punctate, with ern Rivers of New South Wales, and extends shallow punctures (Fig. 28d); vertex weakly as far south as Narooma (Fig. 41). rugopunctate. Pronotum: moderately dense Host plants and biology: Lampromicra punctate; posterior margin of callosite re- aerea is not common in collections, and gion with irregular row of dense punctures. nothing is known of its biology. Single spec- Scutellum: densely punctate, less so on an- imens have been collected on two shrub- terior callus. Thoracic pleura: mostly with like plants (Table 1). dense punctures. Pregenital Abdomen: ster- na densely punctate, lateral regions (laterad Remarks: MCDONALD & CASSIS (1984) in erecting a new synonymy, regarded L. of trichobothria) impunctate. aerea as a geographically isolated colour Vestiture. Body moderately setose. Legs: morph of the ubiquitous L. senator. Our ex- ventral surface of femora, and tibiae densely amination of the same material, re-estab- setose. lishes the species status of L. aerea, on the Structure. Labium: reaching posterior basis of its ground colour (Fig. 28c), red margin of abdominal SIII. Male Genitalia: femora, and colouration of the abdominal pygophore with ventral margin deeply bisin- venter (Fig. 21f), as well as the shape of the uate, with a short medial process; crown of lobal sclerite of CAII(L) (Figs 40c, d). parameres with short hook; ejaculatory appa- Lampromicra regia BERGROTH 1895 ratus well-developed, ventral conducting (Figs 21g, 28d, 41) canal with up to 12 convolutions; CAI ab- sent; CAII bifid; CAII(L) with large, digiti- Lampromicra regia BERGROTH 1895: 287 (n.sp.); DISTANT 1904: 276 (description); SCHOUTEDEN form lobal sclerite, weakly arcute, apex acute; 1904: 30 (list); BERGROTH 1906: 1 (note); FROG- CAII(M) with serrate, hook-shaped lobal GATT 1907: 328 (biology); MCDONALD 1963: 24, sclerite. CAIII fused medially, bifid post-the- 29 (male genitalia); GAEDIKE 1971: 90 (type); cal margin. Female Terminalia: bi-planar, MCDONALD & CASSIS 1984: 557 (description); caudal in orientation; paratergites VIII mod- CASSIS & GROSS 2002: 596 (catalogue) erately-sized, subtriangular; paratergites IX Philia leucochalcea BREDDIN 1903: 58 (n.sp.); small, suboval; gonocoxae I large, medially KIRKALDY 1909: 300 (synonymy) depressed, posterior margin weakly concave. Diagnosis: Lampromicra regia is recog- New specimens examined: Queensland: 1=, nised by the following combination of char- Wongabal S[tate] F[orest] 10 km S Atherton,12 acters: head, most of pronotum, and anteri- June 1992, C Reid, ex beating rainforest bush-

374 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at es/vines (AM); 1=, Upper Nesbit Riv[er], Claudie Creek, Lake Lands, 1500 feet, 16-21 Au- gust 1948, Archibold Expedition, LJ Brass (AMNH). Distribution: Lampromicra regia is re- stricted to tropical north Queensland (Fig. 41). Host plants and biology: Lampromicra regia is known from a few specimens, and there is no recorded host plant. A single species was found in rainforest understorey. Remarks: Body colouration in species of Lampromicra is usually of no diagnostic val- ue, except in species such as L. regia and L. leucocyanea. No other species approximates the colour pattern of the former species, with its yellow and fuscous-iridescent green dorsum. In contrast, its relationships are un- certain, as the genitalia do not provide a ba- sis for sister-group relationships, although they bear a strong resemblance to L. aerea.

Lampromicra senator (FABRICIUS 1803) ta); LETHIERRY & SEVERIN 1893: 48 (incertae Fig. 41: Distribution of Lampromicra (Figs 1d, 21h, 28e, 40, 41) sedis); MCDONALD & CASSIS 1984: 555 (syn- species. Teytra senator FABRICIUS 1803: 131 (n.sp.) onymy) Scutellera dux KIRBY 1818: 474 (n.sp.); STÅL 1873: Callidea collaris WALKER 1867: 40 (n.sp.) DISTANT 15 (synonymy) 1899: 51 (synonymy with Callidea femorata); Scutellera corallifera MACLEAY 1827: 466 (n.sp.); LETHIERRY & SEVERIN 1893: 48 (incertae sedis); STÅL 1873: 15 (synonymy) MCDONALD & CASSIS 1984: 555 (synonymy) Scutellera basalis GRAY 1832: 223 (n.sp.); STÅL Callidea aureocincta WALKER 1867: 40 (n.sp.); 1873: 15 (synonymy) DISTANT 1899: 51 (synonymy with Callidea Tectocoris binotata WESTWOOD 1837: 15 (n.sp.); femorata); MCDONALD & CASSIS 1984: 555 (syn- STÅL 1873: 15 (synonymy) onymy) Calliphara binotata: GERMAR 1839: 129 (new com- Callidea erythrina WALKER 1867: 44 (n.sp.). New bination) Synonymy Callidea senator: GERMAR 1839: 121 (new combi- Tectocoris croesus VOLLENHOVEN 1868: 176 nation); DALLAS 1851: 22 (list); VOLLENHOVEN (n.sp.); STÅL 1873: 15 (synonymy) 1863: 25 (list) Philia fulgurans STÅL 1873: 15 (n.sp.); DISTANT Philia senator: SCHIÖDTE 1842: 284 (new combi- 1888: 476 (note); LETHIERRY &SEVERIN 1893: 22 nation); MAYR 1866: 23 (as Philya senator); STÅL (catalogue); SCHOUTEDEN 1904: 30 (list); MC- 1868: 10 (list); LETHIERRY & SEVERIN 1893: 23 DONALD & CASSIS 1984: 555 (synonymy) (catalogue; synonymy); BREDDIN 1900: 140 (de- Philia senator croesus: TRYON 1982: 14 (New scription); KIRKALDY 1901: 52 (distribution); Guinea; Queensland) SCHOUTEDEN 1904: 30 (list); SCHOUTEDEN 1907: Philia femorata: DISTANT 1899: 35 (new combina- 108 (list) tion); SCHOUTEDEN 1904: 30 (list) Scutellera aurantiacomaculata BLANCHARD 1849: Lampromicra senator: FROGGATT 1902: 318 (new pl. 4 fig. 1 (n.sp.); STÅL 1873: 15 (synonymy); combination); DISTANT 1904: 276 (note); FROG- MCDONALD & CASSIS 1984: 555 (synonymy) GATT 1907: 327 (biology); KIRKALDY 1909: 300 Scutellera metallica MONTROUZIER 1855: 94 (catalogue); HORVÁTH 1919: 307 (description); (n.sp.); STÅL 1873: 15 (synonymy); MCDONALD SZENT-IVANY & CATLEY 1960: 256 (distribution); & CASSIS 1984: 555 (synonymy) MCDONALD 1961: 175-176, 184, figs 5-8 (male Callidea femorata WALKER 1867: 38 (n.sp.); genitalia); MCDONALD &CASSIS 1963b: 232, figs LETHIERRY & SEVERIN 1893: 23 (incertae sedis); 8,9 (female genitalia); MCDONALD 1963c: 277 MCDONALD & CASSIS 1984: 555 (synonymy) (larval morphology); KUMAR 1964: 41 (male gen- Callidea curtula WALKER 1867: 39 (n.sp.); DIS- italia); KUMAR 1965: 41 (morphology); BLACK TANT 1899: 51 (synonymy with Callidea femora- 1968: 573 (distribution); MONTEITH 1982: 534

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(aggregation); JAVAHERY et al. 2000: 492 (biolo- mostly iridescent green, sometimes with gy); CASSIS & GROSS 2002: 596 (catalogue) copper tinge. Philia insignis SCHOUTEDEN 1904: 297 (n.sp.); BERGROTH 1908: 140 (list); MCDONALD & CAS- Texture. Dorsum with dense distribu- SIS 1984: 555 (synonymy) tion of moderately deep punctures; medial Lampromcira senator metallica: SCHOUTEDEN regions of pronotal callosite region and an- 1907a: 108 (type) terior callus of scutellum impunctate (Fig. Philia croesus: SCHOUTEDEN 1907b: 45 (list) 28e). Thoracic pleura: moderate distribu- Lampromicra fulgurans: KIRKALDY 1909: 300 (new tion of deep punctures. Pregenital Ab- combination; catalogue) domen: venter mostly with moderate distri- Lampromicra insignis: KIRKALDY 1909: 300 (new bution of shallow punctures, laterally im- combination; catalogue) punctate (Fig. 28e). Diagnosis: Lampromicra senator is recog- Vestiture. Dorsum and abdominal ven- nised by the following combination of char- ter with dense distribution of elongate, se- acters: dorsum densely setose (Figs 1d, 28e); mi-erect pale setae; thoracic pleura with body mostly iridescent green (Fig. 28e); AI scattered setae (Figs 1d, 28e). Antennae: orange, AII(a)-AIV fuscous; anterior mar- AI-AII(a) sparsely setose; AII(b)-AIV with gin of scutellum most often with broad el- dense distribution of short, semi-erect setae. liptoid callus (usually orange) (Fig. 1d); Legs: ventral surface of femora densely se- femora orange; lateral margins of abdominal tose, with elongate setae; tibiae and tarsi venter impunctate (Fig. 21h); lateral mar- gins of abdominal venter orange, remainder densely setose. iridescent green (Fig. 21h); apex of Structure. Antennae: AI & AII(a) CAII(M) perpendicular and elongate; and, subequal in length; AIV longest segments, a CAIII heavily sclerotized, arcuate. little longer than AIV; AIIII-AIV flattened. Description: Body moderate size, males Labium: reaching posterior margin of ab- 10-12 mm, females 10-13 mm. dominal SIV; LI shortest segment, LII longest segment. Male Genitalia: pygophore Colouration. Body mostly iridescent with broad setal patch on dorsal margin of green to purplish-blue, with orange mark- genital opening; parameres with short hook- ings, sometimes with copper-iridescence shaped apex; phallotheca with moderately (Figs 1d, 28e). Head: uniformly iridescent large subapical thornlike processes; aedea- green; lorum sometimes with copper tinge; gus: ejaculatory apparatus with convoluted gula yellow to orange; antennae, AI orange, ventral conducting canal, 8-10 convolu- AII-AIV fuscous. Labium: ventral surface tions, ejaculatory reservoir elliptoid, dorsal fuscous, dorsal surface mostly orange-brown. conducting canal broad; ductus vesica S- Pronotum: mostly iridescent green, some- shaped, extending beyond conjunctival ap- times with purple tinge; callosite region of- pendages in resting position; CA I absent; ten with copper iridescence (Figs 1d, 28e). CAII(L) with large hook-shaped lobal scle- Thoracic pleura: iridescent green, often rite; CAII(M) distally perpendicular, with with copper-iridescence; peritreme irides- large sub-elliptoid lobal sclerite with basal cent green, evaporative areas matt black. and apical serrations, variable in shape, ta- Thoracic sterna: uniformly fuscous. Scutel- pered to rounded apically; CAIII arcuate, lum: mostly iridescent green, sometimes heavily sclerotised, separated. Female Ter- with purple tinge; anterior callus mostly or- minalia: large, biplanar; paratergites VIII ange, sometimes concolorous with remain- small, subtriangular, medially tapered; der of scutellum (Figs 1d, 28e). Hemelytra: paratergites IX small, subelliptical; gono- endocorium iridescent green; membrane coxae I large, medially swollen; weakly con- black. Legs: coxae, trochanters and femora cave laterally. Spermatheca: fecundation orange; tibiae, tarsi and pretarsi black. Pre- canal elongate; reservoir large, oval; distal genital Abdomen: venter mostly iridescent fecundation canal elongate; proximal and green, with lateral margins orange, with V- distal flanges present; bulb small, elliptoid. shaped inner outline; sometimes venter more orange medially, with iridescent green, Measurements. MCDONALD & CASSIS sublateral band (Fig. 21h). Terminalia: 1984: Table 4.

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Type material examined: Callidea femorata WALK- vember 1979, MS & BJ Moulds, ex sandstone ER: Holotype, ‘Ceram’, ‘B.M. Hem. Type No. 445’ outcrop (AM); Western Australia; 16== (BMNH); Callidea curtula WALKER: Lectotype, Y, 17YY, Kimberley district, Black Rock Pool, ‘Ké’, ‘Wallace’, ‘Saunders 65-13’, ‘B.M. Hem. 18km N of Victoria Hwy, G Cassis and R Silveira, Type No. 444’ (BMNH; here designated); Callidea 24 May 1999, ex Adiantum aethiopicum (AM); collaris WALKER: Lectotype, =, ‘Dor’, ‘Wallace’, 5== 7YY, Kimberley district, Middle Springs, ‘B.M. Hem. Type No. 446’, ‘Saunders 65-13’ 16.6 km N Victoria Hwy, 24 May 1999, G Cassis (BMNH; here designated); Callidea collaris WALK- and R Silveira, ex Ficus leucotricha (AM); 1= ER: Paralectotypes, Y, same data as lectotype 2YY, Beagle Bay, M Granney (AM); 2YY, Je- (BMNH; here designated), =, ‘Ké’, ‘Callidea col- remiah Hills, on Carlton Hill Station, near KJ-8, laris WALK. Cat.’ (BMNH; here designated), =, 14 July 1989, N Poulter (AM); Northern Terri- ‘Cer’ ‘Callidea collaris WALK. Cat.’, ‘Saunders 65- tory: 6== 5YY, Armstrong River, near Top 13’ (BMNH; here designated), 1= 1Y, ‘Ter’ ‘Cal- Springs, 16°41’09“S 132°03’25“E, 9 January lidea collaris WALK. Cat.’, ‘Saunders 65-13’ 2001, MS & BJ Moulds (AM); 3== 1Y, Mann (BMNH; here designated; male pygophore miss- River, Land, 12°28’24“S ing); Callidea aureocincta WALKER: Lectotype, Y, 134°8’58“E, 20 July 1993, P Taçon (AM). ‘Salwatty’, ‘B.M. Hem. Type No. 443’ (BMNH; Distribution: Lampromicra senator is here designated); Callidea erythrina WALKER: Holotype, Y, ‘North Austr.’, ‘Vict. R. Depart.’, broadly distributed in northern Australia, ‘B.M. Hem. Type No. 458’ (BMNH). Callidea cur- and is commonly encountered in tropical re- tula was described from two specimens; from the gions of Queensland, Western Australia and following locations, ‘a. Aru’ and ‘b. Ké’. Only the the Northern Territory. It is also more former specimen was found and has been desig- broadly distributed on the eastern seaboard, nated as the lectotype. Callidea aureocincta was de- extending as far south as Wollongong in scribed from two specimens, as follows: ‘a. New New South Wales (Fig. 41). Guinea. Presented by W.W. Saunders Esq.’ and ‘b. Salwatty. From Mr Wallace’s collection’. The lat- Host plants and biology: Lampromicra ter specimen has been designated as the lectotype. senator is found on many plants (Table 1). A specimen was found that had the following la- MCDONALD (1963c) described aspects of its bel data: ’59-58, Dory, New Guinea’, ‘Callidea col- biology, reporting its primary host as Breynia laris WALK. Cat.’ and ‘Paratype’. In the syntypical oblongifolia. We have found it on a number series of Callidea collaris, there is no mention of a of species of Ficus (Table 1), as well as addi- specimen from New Guinea. We have refrained tional fig species from a recent field trip to from designating this specimen as a paralectotype north Queensland, that are not recorded of either Callidea collaris or C. aureocincta. here. In The Kimberley district of Western Other material examined: Queensland: 1=, Rockhampton, 12 January 1973, MS Moulds Australia, we found this species in aggrega- (AM); 1= 2YY, Flinders Island, 2 June 1923, C tions on a fern (Table 1), at the beginning of Hedley (AM); 3== 1Y Almaden, WD Camp- the dry season, indicating a possible aesti- bell (AM); 1=, Clermont, July 1928, KK Spence vating site. (AM); 1Y 1 larva, Carnarvon Gorge National Remarks: MCDONALD & CASSIS (1984) Park, Track 7-8, 1 December 1993, G Cassis, ex Breynia (AM); 1=, Heathlands, 11°14’S established an extensive synonymy for Lam- 142°35’E, 9 December 1992, P Zborowski & K promicra senator, recognising significant Halfpapp, ex Leptospermum (ANIC); 16== polymorphism in colour and male genitalia. 18YY, Louis Creek, SW Adels Grove, 8 Octo- We confirm in this work, that the condition ber 1963, R Lossin (AM); 1=, 15 km W Paluma, of CAII(M) exhibits significant and contin- A Sundholm & J Bugeja, 16 January 1989; 3== uous variation within and between popula- 2YY, Boolburra, 12 January 1968, G Hangay tions. The apex of CAII(M) is elongate and (AM); 7== 12YY, Foleyvale Arboretum Re- strongly perpendicular to its base, with vari- serve, 20-25 January 1968, G Hangay (AM); ous sclerotizations. The shape of the apex == 5 , Expedition Range, 1 January 1968, G but is not indicative of species limits, as it Hangay (AM); 3== 4YY, Expedition Range, 5 varies from a subtriangular process to being May 1945, TW Gamble (AM); 2== 2YY, Wild Horse Creek, 30 January 1968, G Hangay (AM); rounded apically, and sometimes with sub- 1=, Mt Surprise, 23 February 1988, G O’Reilly apical projections. MCDONALD & CASSIS (AM); New South Wales: 1Y, Mt Ousley, Wol- (1984: Figs 54-58) illustrated these varia- longong, 27 December 1967, RB Lachlan (AM); tions of CAII(M). In the main, other com- 1= 2YY, 30 km W Fairview, NW Laura, 7 No- ponents of the aedeagus are uniform within

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the species. The colour of L. senator also mesosternum sulcate, laterally striate (Fig. varies considerably, although the predomi- 42c); AII(a) arcuate; AII(b) short; antero- nant colour morph is iridescent green, with lateral margin of pronotum concave (Fig. the anterior callus of the scutellum orange. 28b); metathoracic external efferent system Within some populations, the orange scutel- reduced (Figs 42c, d); peritreme short, sub- lar callus is absent, and the scutellum is uni- reniform (Fig. 42c, d); pygophore enlarged, formly iridescent green. There is also varia- medially excavate (Figs 42e, f, 43a); crown tion in the colour of the callosite region (iri- of parameres short (Fig. 43b); CAI absent descent green to iridescent copper) and the (Fig. 43c); CAII tripartite, three elongate pregenital abdominal venter (mostly irides- lobal sclerites (Figs 43c, d); CAIII fused me- cent green with orange lateral margins to dially, apically bifid (Fig. 43d); female ter- mostly orange, with sublateral band of iri- minalia caudally oriented; and, gonocoxae I descent green). Extralimital populations greatly enlarged. have colour patterning not seen in Aus- Description: Large species; elongate- tralian populations. However, the male gen- ovoid (Figs 1e, 28b); dorsally and ventrally italia do not show any discontinuities be- convex; dorsum densely punctate (Figs 1e, tween these populations. 28b); body almost glabrous (Figs 1e, 28b); None-the-less, in considering the extent body iridescent green with orange and black of the morphological polymorphism and markings (Figs 1e, 28b). Head: triangular widespread geographic distribution of L. (Fig. 42a); weakly convex; jugal margins sub- senator, it would be desirous to reinvestigate carinate (Fig. 42b); lorae strongly demarcat- the synonymy of the extralimital junior syn- ed, margins carinate; bucculae narrow, later- onyms (Scutellera iridescenta, Callidea femora- al margins parallel. Antennae: AII(a) ta, Callidea curtula, Callidea collaris, Callidea strongly arcuate, more so in males; AII(b) aureocincta, Tectocoris croesus and Philia ful- very short; AIV flattened. Labium: reaching gurans) using molecular data. In keeping abdominal SIV; LII bicompressed. Prono- with this broad conception of the species, tum: subtrapeziform, shield-like, moderately we have synonymised another of WALKER’s convex (Figs 1e, 28b); post-callosite region names, Callidea erythrina, which has the or- depressed (Figs 1e, 28b); anterior margin ange scutellar callus, and a mostly purple strongly concave (Figs 1e, 28b); anterolater- dorsum, the latter a recurrent artefact of al margins weakly concave, carinate (Figs 1e, preservation. 28b); posterior margin rectilinear (Figs 1e, 28b); post-humeral angles depressed. Scutel- Scutiphora GUÉRIN 1831 lum: elongate, large, covering most of (Figs 1e, 21i, 28b, 42, 43, 44) forewings, base of exocorium and tip of ab- Scutiphora GUÉRIN 1831: pl. XI fig. 7 (gen. nov.); domen visible (Figs 1e, 28b); moderately LAPORTE 1832: 11 (list); GERMAR 1839: 110 (de- convex, strongly declivent posteriorly; ante- scription); AMYOT &SERVILLE 1843: 35 (descrip- rior region raised, with two prominent calli tion); STÅL 1865: 33 (key); MAYR 1866: 16, 23 (Figs 1e, 28b); anterolateral angles upturned. (key, list); STÅL 1873: 8,10 (key, list); LETTHIER- Thoracic pleura: anterior margin of proepis- RY &SEVERIN 1893: 18 (catalogue); SCHOUTEDEN ternum truncate (Fig. 42c); external efferent 1904: 28 (description); KIRKALDY 1909: 308 (cat- system of metathoracic glands reduced, per- alogue); DUPUIS 1952: 452 (taxonomy); MCDON- ALD &CASSIS 1984: 555 (description); CASSIS & itreme short, subreniform, evaporative areas GROSS 2002: 598 (catalogue) confined minimally beyond margins of per- Peltophora BURMEISTER 1835: 393 (gen. nov.); itreme, including mesepimeron (Figs 42c, d). DALLAS 1851: 19 (synonymy); STÅL 1865: 33 Thoracic sterna: prosternum depressed; (key) mesosternum medially sulcate, laterally stri- Type species: Scutiphora: Scutiphora rubromaculata ate; metasternum depressed. Pregenital Ab- GUÉRIN 1831 (= Scutellera pedicellata KIRBY domen: posterolateral angles of abdominal 1826), monotypy; Peltophora: Peltophora cruenta venter rounded; connexiva smooth, without BURMEISTER 1835 (= Scutiphora rubromaculata processes; male SIII medially rounded, with- GUÉRIN 1831), monotypy out medial sulcation or processes; female SI- Diagnosis: Scutiphora is recognised by II with very weak medial depression; male the following combination of characters: SVII visible, weakly overlapping pygophore;

378 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at female SVII recessed, not visible. Male Gen- 7 (n.sp.); LAPORTE 1832: pl. 55 fig. 3 (habitus); italia: pygophore greatly enlarged; posterior GUÉRIN-MÉNEVILLE 1838: 165 (description); margin of pygophore with medial notch (Figs STÅL 1873: 10 (synonymy) 42e, f, 43a); dorsal margin with setal patches Peltophora cruenta BURMEISTER 1835: 393 (n.sp.); (Fig. 42f, 43a); parameres with thick stem, STÅL 1873: 10 (synonymy) Peltophora rubromaculata: HERRICH-SCHAEFFER crown small, weakly notched Fig. 43b); duc- 1836: 100, pl. 10, fig. 326 (new combination); tus seminis proximalis narrow, membrane- BURMEISTER 1835: 393 (list); GERMAR 1839: 110 ous, singular (Figs 43c, d); ventral conduct- (list) ing canal, with up to 12 convolutions (Figs Scutiphora picta GUÉRIN-MÉNEVILLE 1838: 165 43c, d); ejaculatory reservoir subelliptoid, (n.sp.); CASSIS & GROSS 2002: 599 (catalogue) moderately-sized (Figs 43c, d); vesica S- Peltophora picta: AMYOT & SERVILLE 1843: 35 shaped; elongate, extending beyond con- (new combination); DALLAS 1851: 19 (descrip- junctival appendages (Figs 43c, d); CAI ab- tion) sent; CAII tripartite, CAII(L) elongate, Peltophora pedicellata: STÅL 1873: 13 (list); sickle-shaped, strongly sclerotized, CAII(M) LETHIERRY &SEVERIN 1893: 21 (catalogue); basally membraneous, apically with two dig- FROGGATT 1897: 104 (description; biology); FROGGATT 1901: 1594 (description; cherry pest); itiform lobal sclerites (Figs 43c, d); CAIII VAN DUZEE 1905: 188 (list); FROGGATT 1907: 327 strongly sclerotized, fused medially, apically (description; cherry pest) bifid (Figs 43c, d). Female Terminalia: cau- Scutiphora pedicellata:SCHOUTEDEN 1904: 28 (new dally oriented; paratergites VIII moderately combination); KIRKALDY 1909: 291 (catalogue); large, subelliptoid, medially incrassate; TILLYARD 1926: 149 (diagnosis); MCKEOWN 1942: paratergites IX small, suboval, medial mar- 86 (biology); MCDONALD 1961: 176-177, 185 figs gins rounded; gonocoxae I greatly large, sub- 9-13 (male genitalia); MCDONALD 1963a: 30 triangular, dorsal margin excavate; Sper- (male genitalia); MCDONALD 1963b: 231-232, matheca: fecundation canal short; reservoir figs 6,7 (female genitalia); MCDONALD 1963c: round; proximal and distal flanges present; 293, figs 11-16 (life cycle); KUMAR 1964: 60 bulb oval. (male genitalia); KUMAR 1965: 41 (male geni- talia); HELY et al. 1982: 124, 145, 165 (biology; Diversity and distribution: Scutiphora is pest status); MCDONALD &CASSIS 1984: 522, figs a monotypic genus endemic to Australia. 49,50 (description); CASSIS & GROSS 2002: 593 (catalogue) Included species: S. pedicellata (KIRBY 1826) Australia Diagnosis: Scutiphora pedicellata is recog- nised by the following combination of char- Remarks: The affinities of Scutiphora are acters: iridescent green with orange and fus- difficult to deduce, with distinctive charac- cous markings (Figs 1e, 28b); AII(a) arcu- ter states of the antennae, mesosternum, ex- ate; AII(b) very short; labium reaching ab- ternal efferent system of the metathoracic dominal SIII; pronotum with deep trans- glands, and male aedeagus. The metatho- verse furrow (Figs 1e, 28b); scutellum with racic peritreme resembles those found in pair of submedial calli (Figs 1e, 28b); Calliphara and Lampromica, with all three mesosternum striate; abdominal venter genera having the peritreme subreniform in mostly orange, with SII/III fuscous, and SIV- shape, although it is much smaller in Scu- SVII with sublateral iridescent green band tiphora. The male genitalia are autapomor- of markings (Fig. 21i); and, male (Figs 43a- phic, with the CAII tripartite and enlarged d) and female genitalia as in generic diagno- digitiform lobal sclerites. Scutiphora also has sis. fused, U-shaped CAIII, which is not found Description: Large species, males 13-15 in the former two genera, but in genera such mm, 12-15 mm. as Calliscyta and Cantao. Colouration. Dorsum iridescent green Scutiphora pedicellata (KIRBY 1826) with orange and fuscous markings (Figs 1e, (Figs 1e, 21i, 28b, 42, 43, 44) 28b). Head: mostly iridescent green, with Iridescent Jewel Bug anteclypeus and ocular regions fuscous; an-

Scutellera pedicellata KIRBY 1826: 517 (n.sp.); tennae mostly fuscous, AI sometimes with Scutiphora rubromaculata GUÉRIN 1831: pl. XI Fig. green iridescent tinge, AII(a) greatly elon-

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gate, AII(b) short. Pronotum: anterior and BJ Day (AM); 1Y, Clarence, 21 January 1990, R anterolateral margins orange, callosite re- de Keyzer & GA Clark, 21 January 1990, ex Lep- gion mostly fuscous, medially orange, disc tospermum; Australian Capital Territory: Can- uniformly iridescent green. Scutellum: iri- berra (Farrer), 35.22S 149.05E, DCF Rentz, 3-cii- descent green with four to six elliptical fus- 1986 (ANIC); Tasmania: Aspley Gorge, 23 Feb- cous spots. Legs: coxae, trochanters and ruary 1994, J & A Leask (AM). proximal 2/3 of femora orange, remainder of Distribution: Scutiphora pedicellata is femora and tibiae iridescent green, tarsi fus- broadly distributed in New South Wales, ex- cous. Pregenital Abdomen: venter mostly tending as far north as southeast Queens- orange, with submarginal iridescent green land and as far south as Tasmania (Fig. 44). markings (Fig. 21i). Terminalia: uniformly This species was erroneously recorded from orange. New Zealand (CASSIS & GROSS 2002, also references therein). Texture. Iridescent green regions of body densely punctate; orange regions most- Host plants and biology: Scutiphora pedi- ly impunctate (Figs 1e, 28b). cellata is known from five host plants (Table 1). MCDONALD (1963c) described its biolo- Vestiture. Body mostly glabrous (Figs gy and found it feeding primarily on Dodon- 1e, 28b). Antennae: AI & AII(a) glabrous, aea triquetra. AII(b)-AIV setose. Legs: ventral surface of femora weakly spinose; ventral surface of Remarks: This species is remarkably ho- tibiae densely setose, with short thick setae. mogenous, with colour patterning and male genitalia exhibiting little intraspecific varia- Structure. Antennae: AII(a) arcuate, tion. It is easily distinguished by the irides- strongly in male; AII(b) longest segment; cent green body with orange markings (Figs AII(b) very short; AIV little longer than 1e, 28b) and the male genitalia. AIII. Labium: reaching abdominal SIII. Pronotum: anterolateral margins very weak- Subfamily Tectocorinae ly excavate; post-callosite region with deep Tectocorinae MCDONALD & CASSIS 1984: 566 transverse furrow. Scutellum: with pro- (new subfamily); CASSIS & GROSS 2002: 600 nounced, submedial calli on anterior mar- (catalogue) gin. Male (Figs 43a-d) and female genitalia as in generic description. Diagnosis: The Tectocorinae are recog- nised by the following combination of char- Measurements. MCDONALD & CASSIS acters: usually large species (Figs 45a, b); 1984: Table 3. aposematic colouration; elongate-ovoid Material examined: New South Wales: 1Y, (Figs 1f, 45a, b), shallowly punctate; elon- Deua River National Park, near Woolli Home- gate head (Fig. 46a); thoracic sterna flat stead, G Cassis, 9 October 1988 (AM); 1=, Col- (Fig. 46c); anterior margin of proepisternum Y laroy, 28 December 1981, BJ Day (AM); 1 , linear (Fig. 46c); efferent system of Frenchs Forest, La Sorlie Road, December 1987, metathoracic glands vestigial, with rectilin- T Ralph (AM); 1Y, Cowan Field Station, 25-v- ear peritreme (Fig. 46d); males with abdom- 1981, M McDonald (AM); 2YY, Ingleburn, 8 September 1985, R Bejsak (AM); 15YY, Avoca inal androconial glands on SIV-SVI (Figs Beach, 15 November 1985, S Hunter & AL 2g-i, 45c); pregenital abdomen in both sex- Johnson (AM); 5==, NSW: Pearl Beach, 1-i- es without stridulatory vittae; ventral sur- 1986, C Urquhart, ex ground (AM); 2== 1Y, face of pygophore ventrally oriented (e.g., Woodford, Blue Mountains, 30 November 1985, Figs 46c); parameres large, strongly hook- M Dingley (AM); 1= 5YY, Mt Kaputar, 6-xii- shaped (Fig. 47b); ductus seminis proximalis 1979, DJ Scambler (AM); 1= 2YY, Mt Kaputar, broad, bounded by ligamentary tubule (Figs Bullawa Creek, 27 November 1984, G Hangay 47c, d); ejaculatory apparatus box-like, (AM); 5== 28YY, Mt Kaputar, 1500 m, 29 No- heavily sclerotised; without convoluted vember 1984, G Hangay (AM); Whiskers, 7 km ventral conducting canal (Figs 47c, d); three WNW Hoskinstown, 23.241S 149.23E, 18 Febru- ary 1990, MS Upton, ex privet bush (ANIC); conjunctival appendages present, mostly MT Goonoo State Forest, southside, 9 November membraneous (Figs 47c, d); CAII undivided 1987, DK McAlpine & R de Keyzer (AM); 1Y, with conical lobal sclerite (Figs 47c, d); fe- Mann River, near Grafton, 30 December 1978, males lacking interlocking rami; spermathe-

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Fig. 42: Scanning electron micrographs of key characters of Scutiphora pedicelllata (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Ea = evaporative areas; GO = genital opening; Pe = peritreme; P(r) = right paramere. Scale bars = 1 mm.

cal fecundation canal short; spermathecal lowing previous workers (SCHOUTEDEN reservoir membraneous, oval; spermathecal 1904; KIRKALDY 1909; MCDONALD 1961, pump minute, without flanges. 1963b). In our view, this is not supported as Remarks: Tectocoris diophthalmus is a dis- Tectocoris lacks the scutellerine-type ejacu- tinctive taxon based on the male and female latory apparatus. In our view there is suffi- genitalia alone. The aedeagus has a liga- cient evidence to retain the use of the Tec- mentary tubule (Figs 47c, d), which is also tocorinae as a subfamily, pending a more de- found in some species of Pachycorinae, in- tailed cladistic analysis of the family. cluding Agonosoma trilineatum (Fig. 18c). In contrast, Tectocoris has androconial glands Tectocoris HAHN 1834 which are only found in the Odontotarsi- (Figs 1f, 2g-i, 45, 46, 47, 48) nae. SCHUH & SLATER (1995) amongst oth- Tectocoris HAHN 1834: 33 (gen. nov.); DALLAS ers have ignored the use of this subfamily, 1851: 3 (description); STÅL 1865: 33 (key); MAYR placing Tectocoris in the Scutellerinae, fol- 1866: 17 (list); STÅL 1873: 11 (list); LETHIERRY &

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Fig. 43: Male genitalia of Scutiphora pedicellata (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbreviations: C = crown of right paramere; CAII(L) = second conjunctival appendage, lateral branch; CAII(M) = second conjunctival appendage, medial branch; CAIII = third conjunctival appendage; DS(D) = ductus seminis dorsalis; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; SP = setose patch; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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SEVERIN 1893: 19 (catalogue); SCHOUTEDEN 1904: 19 (description); KIRKALDY 1909: 306 (cat- alogue); MCDONALD &CASSIS 1984: 568 (list); CASSIS & GROSS 2002: 600 (catalogue) Type species: Tetyra cyanipes FABRICIUS 1803 (= Cimex diophthalmus THUNBERG 1783), monotypy Diagnosis: Tectocoris is recognised by the following combination of characters: large, elongate-ovoid body (Figs 1f, 45a,b); head elongate (Fig. 46a); anterolateral margins of pronotum strongly divergent posteriorly (Figs 1f, 45a,b); posterior margin of prono- tum excavate (Figs 1f, 45a,b); anterior mar- gin of proepisternum entire (Fig. 46b); ex- ternal efferent system of metathoracic glands reduced, linear (Fig. 46d); mesepimeron without evaporative areas (Fig. 46d); abdominal SIII-SIV sulcate; males with androconial glands on abdomi- nal SIV-VI (Figs 2g-I, 45c); male SVIII vis- ible (Fig. 46e); parameres strongly hooked (Fig. 47b); aedeagus with three conjunctival appendages (Figs 47c,d); CAI medially fused Fig. 44: Distribution of Scutiphora (Figs 47c,d); ductus seminis proximalis clavus visible (Figs 1f, 45a,b); tip of mem- pedicellata. bounded by ligamentary tubule (Figs 47c,d); brane wing visible (Figs 1f, 45a,b). Hemely- ejaculatory apparatus box-like (Fig. 47c), tra: exocorium expanded (Figs 1f, 45a,b). heavily sclerotized; spermathecal reservoir Thoracic Pleura: anterior edge of proepister- greatly enlarged, oval; and, spermathecal num linear, explanate (Fig. 46c); external ef- pump minute. ferent system of metathoracic glands greatly reduced, ostiole minute, peritreme as linear Description: Body elongate-ovoid (Figs sulcation (Fig. 46d); evaporative areas re- 1f, 45a,b); aposematic colouration, often duced, not extending to mesepimeron (Fig. sexually dimorphic (Figs 1f, 45a,b), females 46d). Thoracic Sterna: prosternum de- mostly oranges, dorsum often with iridescent pressed (Fig. 46c); mesosternum weakly im- green or blue markings (Figs 1f, 45a,b), pressed, laterally striate; metasternum flat. males ranging from uniformly orange to Pregenital Abdomen: laterally rounded, pos- mostly iridescent green or blue, with orange terior angles weakly acuminate (Fig. 45c); markings (Figs 1f, 45a,b); dorsum densely abdominal SIII-SIV medially sulcate, hous- punctulate (Figs 1f, 45a,b); body nearly ing labium; males with androconial glands glabrous (Figs 1f, 45a,b). Head: elongate, lit- tle longer than wide, porrect, weakly convex present sublaterally on SIV-VI (Figs 2g-i, (Fig. 46a); lateral margins of jugae rectilin- 45c); male SVIII visible, strongly overlap- ear, carinate in profile (Fig. 46a); clypeus ex- ping posterior margin of pygophore (Fig. tending beyond jugae (Fig. 46a); bucculae 46e). Male Genitalia: posterior region of py- elongate, margins low (Fig. 46a); eyes small. gophore visible, margin notched medially Pronotum: callosite region and disc co-pla- (Figs 46e,f, 47a); genital opening broad, nar, without transverse furrow (Figs 1f, densely setose (Figs 46e,f, 47a); parameres 45a,b); anterolateral margins strongly diver- with arcuate, broad stem, densely setose, gent, weakly sinuate, explanate (Figs 1f, crown strongly hooked (Fig. 47b); aedeagus 45a,b), carinate in profile (Fig. 46b); humer- large; phallotheca sub-cylindrical, without al angles angulate (Figs 1f, 45a,b); posterior processes (Figs 47c,d); ductus seminis proxi- margin excavate (Figs 1f, 45a,b). Scutellum: malis narrow, bounded by ligamentary-de- strongly convex; strongly declivent beyond rived membraneous tubule (Figs 47c,d); CAI midpoint (Figs 1f, 45a,b); lateral margins large, lobe-like, without sclerotization (Figs convex (Figs 1f, 45a,b); base of corium and 47c,d); CAII membraneous lobe, with short,

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subfamily Scutellerinae. MCDONALD (1961, 1963b) also placed it within the tribe Scutellerini, but proposed that it occupies a taxonomic position of its own. KUMAR (1965) placed the species in the tribe Pachycorinae, a group otherwise absent from the Australian continent.

Tectocoris diophthalmus (THUNBERG 1783) (Figs 1f, 2g-i, 45, 46, 47, 48) Cotton Harlequin Bug

Cimex lineola FABRICIUS 1781: 340 (n.sp.; junior homonym of Cimex lineola LINNAEUS 1758); FABRICIUS 1787: 281 (description); FABRICIUS 1794: 84 (description); VOLLENHOVEN 1863: 8 (synonymy); STÅL 1873: 19 (synonymy); LETHIERRY & SEVERIN 1893: 19 (synonymy) Cimex diophthalmus THUNBERG 1783: 30, pl. 2 fig. 45 (n.sp.) acute lobal sclerite (Figs 47c,d); CAIII large, Tetyra lineola: FABRICIUS 1803: 135 (new combi- membraneous, medially sclerotized, fused nation) basally, with subapical sclerotized thorn-like Tetyra cyanipes FABRICIUS 1803: 133 (n.sp.); process (Figs 47c,d); vesica moderate size, WOLFF 1811: 171, fig. 165; BLANCHARD 1840: subequal to uninflated CAII, S-shaped (Figs 159 (synonymy); LETHIERRY & SEVERIN 1893: 19 47c,d). Female Terminalia: paratergites VIII (synonymy) moderate size, triangular, medially not con- Cimex banksii DONOVAN 1805: 29 (n.sp.); DALLAS tiguous in ventral view; paratergites IX small 1851: 16 (synonymy); VOLLENHOVEN 1863: 8 (synonymy); STÅL 1873: 19 (synonymy); suboval, medially separated; gonocoxae I LETHIERRY & SEVERIN 1893: 19 (synonymy) moderate size, subelliptoid, medially con- Fig. 45: Habitus and abdominal venter of Scutellera schoenherri ESCHSCHOLTZ 1822: 99 Tectocoris diophthalmus (a) = (b) Y (c) tiguous. Spermatheca: fecundation canal (n.sp.); BURMEISTER 1835: 396 (list); GERMAR abdominal venter. moderate size; reservoir large, oval, membra- 1839: 133 (synonymy); VOLLENHOVEN 1863: 8 neous; spermathecal pump minute, with (synonymy); STÅL 1873: 19 (synonymy); flanges vestigial, pump oval with acute apex. LETHIERRY & SEVERIN 1893: 19 (synonymy) Diversity and distribution: Tectocoris is a Scutellera banksii: GUÉRIN-MÉNEVILLE 1838: 1555 monotypic genus which is known primarily (new combination; description); GERMAR 1839: 133 (synonymy); HERRICH-SCHAEFFER 1839: 1 from the Melanesian and Australian subre- (description); AMYOT &SERVILLE 1843: 28, pl. 1 gions of the Australian zoogeographic re- fig. 5 (description; synonymy); MONTROUZIER gion, as well as Indonesia. 1855: 91 (); WESTRING 1858: 50 Included species: (); MONTROUZIER 1858: 243 (New T. diophthalmus (THUNBERG 1783) Caledonia) Australia, Indonesia, Tectocoris cyanipes: HAHN 1834: 34 (new combi- New Caledonia, Papua New Guinea, nation); DALLAS 1851: 16 (synonymy); VOLLEN- Samoa, Tonga, Vanuatu HOVEN 1863: 8 (description; synonymy; varieties) Tectocoris diophthalmus: HAHN 1834: 33 (new Remarks: The taxonomic history of this combination); DALLAS 1851: 16 (synonymy); MAYR 1866: 22 (synonymy); STÅL 1871: 617 genus and species is convoluted, owing in (synonymy); DISTANT 1920: 144 (New Caledo- part to the extreme colour variability found. nia); SIMMONDS 1922: 36-38 (cotton pest; om- This species has been placed in a number of nivory); BALLARD 1925: 542 (pest status); BAL- genera with varying specific epithets and LARD & HOLDAWAY 1926: 329 (biology); TILL- subspecies. A list of these synonyms can be YARD 1926: 149, pl. 12 fig. 13 (diagnosis; habi- found in CASSIS & GROSS (2002) and tus); BALLARD 1927: 604 (cotton pest); SIM- above. SCHOUTEDEN (1904) and KIRKALDY MONDS 1928: 10-12 (cotton pest; Fiji); DAMMER- (1909) placed the insect in the tribe Scutel- MAN 1929: 2020 (biology); SCHOUTEDEN 1933: 47 lerini with other members of the current (distribution); MCKEOWN 1933: 24 (maternal

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Fig. 46: Scanning electron micrographs of key characters of Tectocoris diophthalmus (a) Head, dorsal view (b) Head, lateral view (c) Head and thorax, ventral view (d) External efferent system of metathoracic glands, ventral view (e) male pygophore, ventral view (f) male pygophore, dorsal view. Abbreviations: Ea = evaporative areas; Pe = peritreme; P(r) = right paramere; SVIII = eighth abdominal sternite. Scale bars = 1 mm.

care); PENDERGRAST 1957: 22 (spermatheca); MANN & STEINBAUER (2001): 12 (); SZENT-IVANY & CATLEY 1960: 256 (host plant; MILLAR 2005: 78 (glandular chemistry); MONTEI- New Guinea); MCDONALD 1961: 177 (male gen- TH 2006: 1135-1152 (maternal care) italia); MCDONALD 1963a: 30 (male genitalia); Scutellera cyanipes: BURMEISTER 1835: 396 (new MCDONALD 1963b: 233, 236 (female genitalia); combination); AMYOT & SERVILLE 1843: 40, 94, MCDONALD 1963c: 289-290, figs 29-31 (life cy- pl. 9 fig 58, pl. 24 fig. 167 (description; synonymy) cle); KUMAR 1964: 43, 49 (male genitalia); Scutellera cyanipoda BOISDUVAL 1835: 624 (n.sp.); BLACK 1968: 574 (distribution); SMITH 1978: VOLLENHOVEN 1863: 8 (synonymy); STÅL 1873: 821-822 (gland chemistry); WILSON et al. 1983: 19 (synonymy); STÅL 1873: 19 (synonymy); 311-317 (biology); CARAYON 1984: 113-134 (an- LETHIERRY & SEVERIN 1893: 19 (synonymy) droconial glands); MCDONALD & CASSIS 1984: Scutellera tongae BOISDUVAL 1835: 624 (n.sp.); 568, figs 78-80 (synonymy; morphology); STAD- BLANCHARD 1840: 159 (synonymy); VOLLEN- DON et al. 1987: 227-234 (gland chemistry; mor- HOVEN 1863: 8 (synonymy); STÅL 1873: 19 (syn- phology); JAVAHERY et al. 2000: 491 (biology); onymy); LETHIERRY & SEVERIN 1893: 19 (syn- CASSIS & GROSS 2002: 603 (catalogue); NAU- onymy)

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Fig. 47: Male genitalia of Tectocoris diophthalmus (a) pygophore, dorsal view (b) paramere, lateral view (c) aedeagus, lateral view (d) aedeagus, ventral view. Abbre- viations: C = crown of right paramere; CAI = first conjunctival appendage; CAII = second conjunctival appendage; CAIII = third conjunctival appendage; DS(P) = ductus seminis proximalis; Er = ejaculatory reservoir; GO = genital opening; Pt = phallotheca; S = stem of paramere; Sg = secondary gonopore; V = vesica; VCC = ventral conducting canal of ejaculatory apparatus; VM = ventral margin of genital opening.

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Tectocoris gambiae WESTWOOD 1837: 14 (n.sp.); STÅL 1873: 19 (synonymy); LETHIERRY & SEV- ERIN 1893: 19 (synonymy) Pachycoris lineola: GERMAR 1839: 133 (new com- bination) Scutellera cyanipes: AMYOT & SERVILLE 1843: 28 (new combination) Tectocoris lineola: DALLAS 1851: 16 (new combi- nation); STÅL 1873: 19 (synonymy); LETHIERRY & SEVERIN 1893: 19 (synonymy); DISTANT 1899: 33 (synonymy); TILLYARD 1926: 149 (diagnosis); MCKEOWN 1942: 83 (biology); HORI 2000: 17 (biology) Tectocoris banksii: DALLAS 1851: 16 (new combi- nation); WALKER 1867: 12 (synonymy) Tectocoris cyanipes: DALLAS 1851: 16 (new combi- nation); VOLLENHOVEN 1863: 8 (description; va- rieties); WALKER 1867: 12 (distribution) Tectocoris bancksii: MONTROUZIER 1861: 60 (in- correct subsequent spelling) Tectocoris obliquus WALKER 1867: 13 (n.sp.); LETHIERRY & SEVERIN 1893: 19 (synonymy) Tectocoris pusillus WALKER 1867: 13 (n.sp.); LETHIERRY & SEVERIN 1893: 19 (synonymy) Tectocoris amboinensis WALKER 1867: 14 (n.sp.); LETHIERRY & SEVERIN 1893: 19 (synonymy) with iridescent blue-green markings (Figs 1f, Fig. 48: Distribution of Tectocoris Tectocoris diophthalmus rufus STÅL 1871: 617 diophthalmus. (n.ssp.) 45a,b); dorsum of females mostly orange, Tectocoris diophthalmus tagalicus STÅL 1871: 617 with iridescent blue-green markings, some- (n.ssp.) times uniformly orange (Figs 1f, 45a,b). Tectocoris diophthalmus schoenherri: STÅL 1871: Head: males orange to mostly iridescent 617 (subsp. arrangement) blue-green, sometimes with purplish hue; fe- Tectocoris lineola banksi: DODD 1904: 483 (mater- males either uniformly orange; bicoloured nal care) morphs with clypeus orange, remainder iri- Tectocoris diophthalmus cookiana KIRKALDY 1909: descent blue-green; underside of head uni- 306 (unnecessary nom. nov. for Tetyra cyanipes: formly orange, sometimes with iridescent BLANCHARD 1841) Tectocoris diophthalmus venusta KIRKALDY 1909: blue-green markings, or uniformly iridescent 307 (unnecessary nom. nov. for Scutellera cyanipo- blue-green. Antennae: mostly fuscous, da var.) sometimes with iridescent green tinge. Labi- Tectocoris purpureus: KNIGHT et al. 1985: 851-853 um: either uniformly fuscous, or in pale (incorrect subsequent spelling; glandular chem- morphs with LI-LII orange, remainder fus- istry) cous. Pronotum: males orange to mostly iri- Diagnosis: Tectocoris diophthalmus is descent blue-green, sometimes in darker recognised by the following combination of morphs with orange spot at midline of cal- characters: male dorsum mostly iridescent losite region; females either orange, or bi- blue-green, with orange markings (Figs 1f, coloured, mostly orange, with anterolateral 45a,b); females mostly orange, with irides- margins iridescent blue-green, with pair of cent blue-green markings (Figs 1f, 45a,b); large submedial iridescent blue-green, some- AIV longest segment; labium reaching ab- times also with smaller sublateral blue-green dominal SIV; male (Figs 47a-d)and female markings. Scutellum: males orange to most- genitalia as in generic diagnosis. ly iridescent blue-green, with purplish hue, intermixed with orange markings anterolat- Description: Large species, males 13-18 erally, mediolaterally, and subdistally, some- mm, females 16-22 mm. times markings coallesced. Hemelytra: exo- Colouration. Dorsum of males mostly corium orange to iridescent blue, with pur- iridescent blue-green, with orange mark- ple hue. Thoracic Sterna: orange to orange ings, sometimes orange, or mostly orange, with red-fuscous, sometimes with green iri-

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descence. Thoracic Pleura: orange to irides- Bejsak (AM); 1Y, Mosman, Sydney, 5 August cent blue-green, always with anterior mar- 1989, McBride (AM); Northern Territory: 1=, gin of proepisternum orange; supracoxal East Point Reserve Lookout, 10 March 1997, M YY lobes most often orange. Legs: femora often Hoskins, ex Hibiscus tiliaceus (AM); 2 , Dar- win, 14 December 1994, A Keast (AM); New orange, sometimes fuscous-iridescent blue Caledonia: 1=, Noumea, 2-iii-1982, DJ Scambler distally; tibiae and tarsi fuscous-iridescent (AM); Philippines: 3== 8YY, Marindupue Is- blue with purplish hue. Pregenital Ab- land, February-March 1991, L Layron (AM). domen: either uniformly orange (Fig. 45c) to red-orange, or orange to red-orange with Distribution: This species is broadly dis- posterior 3/5 of lateral regions of SIII to SIV tributed in Australia from tropical north or SVI iridescent blue-green; male SVIII or- Queensland (including the Torres Strait Is- ange, sometimes with iridescent blue hue. lands) to southeast New South Wales (in- Female Terminalia: uniformly orange to cluding ). It is also found mostly iridescent blue-green. to the west in the Northern Territory (Fig 48). A single specimen was collected in Texture. Dorsum densely punctate with metropolitan Perth. This species is found shallow punctures, ventral surface impunc- extralimitally in the southwest Pacific, and tate (Figs 1f, 45a,b). is also known from the Oriental region (In- Vestiture. Antennae: AI-AII weakly se- donesia). tose, AIII-AIV more densely setose with very Host plants and biology: The biology of short setae. Legs: femora with scattered short Tectocoris diophthalmus has been described by setae, ventral surface of tibiae with more various authors (DODD 1904; MCDONALD dense short setae. Pygophore: genital open- 1963c; SMITH 1978); WILSON et al. 1983; ing with dense distribution of yellow setae. STADDON et al. 1987; MONTEITH 2006). It Structure. Antennae: AII(a) just short- occurs on a wide range of malvaceous er than AI; AII(b) and AIV subequal in plants, including Hibiscus and Malva length; AIII longest segment. Labium: species, Lagunaria patersonia, and cotton, on reaching between abdominal SIV to SV; LII which it is known as a pest. We have found longest segment; LI, LIII and LIV roughly this species on L. patersonia on Lord Howe subequal in length; male and female geni- Island, where it is endemic (also on Norfolk talia as in generic description. Island). This plant has also been introduced Type material examined: Tectocoris pusillus WALK- to eastern Australia as a parkland tree, and ER: Holotype, =, ‘New Caled’, ‘B.M. Hem Type in Sydney, T. diophthalmus is prolific on it in No. 411’ (BMNH); Tectocoris obliquus WALKER: late spring and summer. Its primary host in = Holotype, , ‘56-85’ ‘B.M. Hem Type No. 412’ the northern half of eastern coastal Aus- (BMNH). The type of Tectocoris ambionensis tralia is beach hibiscus, Hibiscus tiliaceus, as WALKER is destroyed. well as on island of Indonesia and southwest Other material examined: Queensland: 1Y, Newell Beach, 16°25’25“S 145°24’22“E, 29 April Pacific. It is also found in suburban gardens 1998, G Cassis, Site Q98-22 (AM); 1Y, Macgre- on Hibiscus cultivars along the eastern gor, 30 August 1988, CE Chadwick (AM); seaboard of Australia. The native distribu- 2YY, Burrum Heads, 14-viii-1982, NW Rodd tion of Tectocoris diophthalmus is unknown, (AM); 1Y, Black Rock, N Townsville, 18 June but distribution records, indicate it has 1991, T Woodger (AM); 2== 2YY, Torres spread further south in Australia over the Strait Islands, Moa Island, 20 February 1975, E past 50 , possibly tracking plantings of Cameron, (AM): 1Y Torres Strait Islands, malvaceous plants. Monteith (pers. comm.) Prince of Wales Island, 16 February 1975, E Cameron (AM); 11== 11YY, Sydney, 11 has found this species commonly on Brachy- March 1992, G Hangay (AM); 1=, Alstonville, chiton acerifolius (Malvaceae [formerly Ster- April-May 1991, B Turner (AM); 2YY, Double culiaceae]) in the Brisbane region, and on Bay, Sydney, 4 February 1999, C Lemann (AM); the inland “bottle trees“, B. australis and B. 2YY, Avoca Beach, 24 February 1985, S Hunter rupestris in the Roma district. (AM); 8YY, Hazelbrook, 1984, M Dingley (AM); 1=, Bundjalong National Park, 32.24S Remarks: Tectocoris diophthalmus is the 152.32E, 15 November 1993, G Cassis, ex beach most emblematic of all the Australian jewel wash (AM); 1Y, Ingleburn, 8 September 1985, R bugs, particularly in eastern Australia where

388 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at it is commonly encountered. Its colour vari- with molecular data required to test its cur- ation is one of the most extreme of any true rent polymorphic definition presented in bug known to us; ranging from orange to this work. metallic blue. It has a complex synonymy The major taxonomic issue in the that has not been re-examined in modern scutellerids worldwide is the classification times. The intrapopulation variation in impediment (sensu CASSIS et al. in press), at colour (e.g., Fig. 1f) is an indication that the both the generic and suprageneric levels. In current synonymy is legitimate, and we have this work we have attempted to establish found that the male genitalia are invariant new character definitions of the genera across the distributional range of this found in Australia, but with reference to the species; inside and outside of Australia. variation found in genera with extralimital species. None-the-less, the definition of Incertae Sedis many genera, particularly in the aposematic Sphaerocoris subnotatus WALKER 1868: 505 (n.sp.); Scutellerinae has serious shortcomings. For DISTANT 1899: 52 (type location); CASSIS & example, the generic boundaries of Brachy- GROSS 2002: 603 (catalogue; incertae sedis) aulax, Calliphara, Chrysocoris, Lampromicra Remarks: The identity of this species is and Lamprocoris are in need of serious revi- unknown. MCDONALD & CASSIS (1984) did sion. LYAL (1979) provided some clarity as not refer to it. CASSIS & GROSS (2002) re- to the generic boundaries for Calliphara, and ferred to it as incertae sedis, and indicated we have attempted in this work to differen- that the type was in Museum Victoria. Ken tiate it from Lampromicra. Moreover, the re- Walker (pers. comm.) has indicated that the lationships of the suprageneric groups of type does not occur in that collection. The scutellerids have received little attention. type locality is given in the original descrip- FISCHER (2001) has made an important step tion as ‘N Australia’. We also examined the in this direction, with the first phylogenetic BMNH collection, where most of the Fran- analysis of genera. This work is hampered cis Walker types are deposited, and did not however by the lack of generic definition for find the type specimen. We have not been many scutellerids. The suprageneric classifi- able to provide any additional clarification cation of scutellerids is similarly impeded, as to its identity or whether it is a synonym with the landmark works of STÅL (1872, of an existing species. Consequently, we re- 1873) and LESTON (1952d), being without tain its status as incertae sedis. serious re-assessment. FISCHER (2001) has also addressed this issue but found, as we Discussion have, that the Scutellerinae and Odonto- The Scutelleridae of Australia are de- tarsinae are ‘convenience groups’ and can pauperate, with only 5 % of the world’s fau- only be defined polythetically at this time. na. The family is much more diverse in the CASSIS et al. (in press), amongst others, tropical regions of southeast Asia and South have stated the importance of globally America, where genera with high species di- scoped taxonomic studies for establishing the framework for correct species assign- versity are known (KIRKALDY 1909), with many species remaining to be described, ment. This is true for the Scutelleridae and particularly in the Neotropical region the shortfalls in the present classification is (Grazia pers. comm.). There is little doubt a consequence of its piecemeal and regional that we have now reached a levelling off of development. the species description accumulation curve The generic and species endemicity of for the jewel bugs of Australia, with only Australian scutellerids is 58 % and 68 % re- three new species described in this work. As spectively, which is commensurate for all of with other places in the world (EGER & LAT- the Pentatomomorpha of Australia (CASSIS TIN 1995), there is also significant new syn- & GROSS 2002: 53 % and 83 %), although onymy in this work. However, a caveat it is a little lower for scutellerid species en- needs to be added in this respect, as the con- demicity. Of the genera represented in Aus- specificity of all the populations of Lam- tralia and extralimitally, there are four ma- promicra senator requires closer scrutiny, jor patterns observed, as follows: i. Oriental

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+ Australia (including Melanesia) + Pacific tion of the scanning electron micrographs. (Coleotichus); ii. Oriental + Australia (Cal- Ms Finlay (AM) prepared the illus- liphara and Lampromicra); iii. Indo-Pacific + trations of the male genitalia and the habi- Australia (Cantao); and, iv. Australia + tus of Heissiphara minuta. Gareth Carter Melanesia + Indonesia (west of Wallace‘s (AM) assisted with the preparation of the line) (Tectocoris). These area relationships distribution maps. Dr. David Britton (AM) are consistent with those found for other sourced the habitat photograph of Choero- pentatomomorphans (CASSIS & GROSS coris paganus. Geoff Monteith (Queensland 2002), although there are no Gondwanan Museum) gave invaluable support by sourc- area relationships exhibited. The Afrotropi- ing most of the habitat photographs (Figure cal connection shown by Cantao is reminis- 1), loan of new material of Calliscyta stalii, cent of the Indo-Pacific pattern described by and technical advice about jewel bug natu- SCHUH & STONEDAHL (1986), but with an ral history. Mr Steve Wilson (Queensland Australian extension; a pattern seen in oth- Museum) provided the habitat photographs er true bugs, e.g., the mirid genus Hypsoleo- (Figures 1a, b, d-f). Dr. Ken Whitney (Rice cus (Schuh pers. comm.). The relationships University) generously gave valuable un- of Morbora are of great interest, as they are published information about the biology of not related to any other Australian scutel- Coleotichus costatus. Dr. Tim Heard and Mr lerid, and our observations indicate a closer Richard Chan (CSIRO, ) also relationship with the Palearctic genus kindly provided unpublished information Psacasta. This hypothesis is however con- and specimens of the introduced species founded by the likely non-monophyly of the Agonosoma trilineatum. Mr Tom Weir Odontotarsinae. (CSIRO, Entomology) loaned material and gave valuable advice. Within Australia, scutellerids are broad- ly distributed in Australia (Figs 6, 15, 23, 27, Dr. Randall Schuh (American Museum 34, 41, 44 and 48), and we give significant of Natural History) is thanked for his col- range extensions for many of the species laboration with the senior author over the (e.g., Coleotichus artensis). We have also past ten years, in undertaking a systematic found distribution patterns that are also survey of Heteroptera across temperate Aus- found in unrelated taxa of true bugs, as fol- tralia, which provided new jewel bug mate- lows: i. restricted to the tropics: e.g., Cal- rial and new host plant information. Ms liphara regalis, Calliscyta stalii, Choerocoris lat- Rossana Silveira (AM) also assisted in this tini and Lampromicra regia (CASSIS et al. fieldwork. GC would particularly like to 2003; Vannius complex [Miridae]); ii. east- thank Jing-Fu Tsai (National Chung Hsing ern Australia: Coleotichus artensis, C. excel- University, Taichung, Taiwan) for detailed lens and Calliphara imperialis (CASSIS & SIL- discussions on the structure of the male VEIRA 2002; alaticollis species-group aedeagus, particularly the ejaculatory appa- []); and, iii. temperate Aus- ratus; his original work has influenced our tralia: Choerocoris grossi, Lampromicra aerea thinking. Dr. Christian Fischer (Freie Uni- and Scutiphora pedicellata (CASSIS & SIL- versität Berlin) is particularly thanked for VEIRA 2001; Nerthra elongata species-group his contribution to this work; pro- [Gelastocoridae]). We have also found that viding information about his own observa- a number of species are ubiquitous across tions on the scutellerids and an 11th hour re- continental Australia (e.g., Coleotichus view of the manuscript. His efforts greatly costatus, Choerocoris paganus and Morbora improved this work. Dr. Jocélia Grazia (Uni- australis). versidade Federal do Rio Grande do Sul, Porte Allegri, Brazil) are also thanked for their generous advice on morphological and Acknowledgements classification issues relating to the Scutel- We especially thank Michael Wall (San leridae. GC would like to thank Paul Brown Diego Natural History Museum) for prepar- and Mick Webb of the Natural History Mu- ing the materials and plate of the key char- seum (London) who generously provided acters of Heissiphara minuta. Ms Sue Lindsay the senior author with free access to the (AM) provided assistance with the prepara- BMNH collection and library, and where

390 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at detailed examination of critical synoptic BER werden mit C. excellens (WALKER) syn- collections were made. GC finished this onymisiert. Callidea erythrina WALKER wird manuscript at the Natural History Museum, mit Lampromicra senator synonymisiert. Lec- found new species records for Australia, as totypen werden für Coleotichus testaceus well as correcting misconceptions about the WALKER, C. excellens, Sphaerocoris circulife- distribution of androconial glands in the rus (WALKER), Callidea aureocincta WALKER, Scutelleridae. GC would like to thank Dr. Callidea collaris WALKER und Callidea curtula Michael Schwartz for lending his laptop in WALKER festgelegt. Die Klassifikation der London, when the senior author’s laptop Scutelleridae wird diskutiert und Beschrei- collapsed. The senior author would like to bungen der in Australien heimischen thank Drs Greg Edgecombe and Chris Reid Unterfamilien präsentiert. Für the subdista- of the Australian Museum, who provide a le Erweiterung des Ductus seminis, inklusive constant source of encouragement and in- des ventralen Verbindungskanals, des Sper- terest in the senior author’s work. mienreservoirs, und des dorsalen Verbin- dungskanals wird ein neuer Begriff, Ejakula- Finally, we would like to thank Dr. tionsapparat, eingeführt. Ein Überblick der Wolfgang Rabitsch, who not only invited us männlichen Pheromondrüsen der pregenita- to participate in the Heiss Festschrift, but len Abdominalsternite zeigt zwei Typen: also accommodated our lateness, extreme Androconien (in den Odontotarsinae und lateness, in producing this work. Tectocorinae) und behaarte Sternaldrüsen (in den Elvisurinae). Die Biologie der aus- Zusammenfassung tralischen Arten wird zusammengefasst und Alle australischen Gattungen und Ar- neue Wirtspflanzenbindungen für Austroti- ten der Scutelleridae werden beschrieben chus rugosus GROSS, die heimischen Arten und mit je einem Beispiel der männlichen von Coleotichus (C. artensis (MONTROU- Genitalien jeder Gattung illustriert. Ras- ZIER), C. costatus FABRICIUS und C. excel- terelektronenmikroskopische Aufnahmen lens), Calliscyta stalii (VOLLENHOVEN), Scuti- werden für einige Merkmale präsentiert. Die phora pedicellata (KIRBY) und Tectocoris australische Schildwanzenfauna enthält 13 diophthalmus (HAHN) mitgeteilt. Gattungen und 25 Arten. Heissiphara wird als neue Gattung beschrieben, mit der einzi- References gen Art, H. minuta nov.sp. aus dem west- AGARWAL S.B. & H.N. BAIJAL (1984): Morphology of lichen Australien. Calliscyta wird als Gat- male genitalia of Scutellerinae and Pen- tung aus der Synonymie mit Choerocoris er- tatominae (Heteroptera) with remarks on in- hoben. Ein Bestimmungsschlüssel für die ter-relationships within the family Pentatomi- australischen Scutelleridae wird präsentiert. dae. — J. Entomol. Res. 8: 52-60. Zwei neue Arten der Gattung Choerocoris ALDRICH J.R. (1988): Chemical ecology of the Het- werden aus dem östlichen Australien be- eroptera. — Annu. Rev. Entomol. 33: 211-238. schrieben: C. grossi nov.sp. und C. lattini ALDRICH J.R. (1996): Sex pheromones in and Heteroptera. — In: SCHAEFER C.W. (Ed.), nov.sp. Lampromicra aerea (DISTANT) wird Studies on Hemipteran Phylogeny. Entomo- aus der Synonymie mit L. senator (FABRI- logical Society of America, Lanham: 199-233. CIUS) erhoben. Calliphara nobilis (LINNAEUS) AMYOT C.J.B. & A. SERVILLE (1843): Histoire Naturelle wird erstmals aus Australien gemeldet. Cal- des Insectes. Hémiptères. — Fain et Thunot, liphara billardierii (FABRICIUS) und C. prasli- Paris: 1-675 + errata. nia praslinia BREDDIN sind nicht Teil der aus- ATKINSON E.T. (1887): Notes on Indian Rhynchota, tralischen Fauna. Die Identität von Sphaero- Heteroptera, No. 2. — J. Asiat. Soc. Beng. 56 coris subnotatus WALKER ist unbekannt und (2): 145-205. als incertae sedis gewertet. Auch die neotro- BAKER A.D. (1931): A study of the male genitalia of pische Art Agonosoma trilineatum (FABRI- Canadian species of Pentatomidae. — Can. J. Res. 4: 148-220. CIUS), die zur biologischen Schädlingsbe- kämpfung gegen das Unkraut Jatropha gossy- BALOCH G.M., MOHYUDDIN A.I. & M.A. GHANI (1968): pifolia (Euphorbiaceae) eingeführt wurde, Xanthium strumarium L. - Insects and other organisms associated with it in West Pakistan. wird beschrieben. Coleotichus borealis DIS- — Comm. Inst. Biol. Cont. Tech. Bull. 10: 103- TANT und C. (Epicoleotichus) schultzei TAEU- 111.

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HERRICH-SCHAEFFER G.A.W. (1840): Die Wanzenarti- KIRKALDY G.W. (1909): Catalogue of the Hemiptera gen Insecten. Getreu nach der Natur abge- (Heteroptera) Vol. 1. . — Felix bildet und beschrieben. Vol. 5 — C.H. Ze- Dames, Berlin: xl, 1-392. h’sche Buchhandlung, Nürnberg: 61-80. KNIGHT D.W., STADDON B.W. & M.J. THORNE (1985): HILGENDORF J.H. & R.D. GOEDEN (1982): Phy- Presumed sex from androconial tophagous insects reported worldwide from glands of male cotton harlequin bug, Tecto- the noxious weeds spiny clotbur, Xanthium coris purpureus (Heteroptera: Scutelleridae) spinosum, and cocklebur, X. strumarium. — identified as 3,5 dihydroxy-4-pyrone. — Z. Bull. Entomol. Soc. Am. 28 (2):147-152. Naturforsch. 40: 851-853.

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Address of the Authors: Dr. Gerry CASSIS BSc Loren VANAGS Australian Museum 6 College Street Sydney Australia E-Mail: [email protected]

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