Bioacoustics of Scutelleridae

Key to Argentinean Scutelleridae 1- Stridulatory areas present on each side of abdominal venter; antennae five-segmented....... ........................... Pachycorinae .....2 - Stridulatory areas absent on abdominal venter; antennae three-segmented........................... .................. Scutellerinae ....... Augocoris 2- Distance from ostiole to metacoxae shorter than distance from ostiole to lateral margin of metapleuron.........................................3 - Distance from ostiole to metacoxae longer than or subequal to distance from ostiole to lateral margin of metapleuron..............................10 3- Ostiole attended by ruga or sulcus that is at least three times as long as wide.............4 - Ostiole not attended by ruga or sulcus or attending structure less than twice as long as wide............8 4- Ostiolar rugae reaching anterolateral corner of metathoracic evaporative area..........5 - Ostiolar rugae not reaching anterolateral corner of metathoracic evaporative area.................6 5- Anterolateral pronotal margins straight or slightly sinuous........................... Lobothyreus - Anterolateral pronotal margins concave............ ................................................. Crathis (*) 6- Ostiolar rugae strongly curved anterad; male genital cup and female basal plates concealed by last abdominal sternite............ Agonosoma - Ostiolar rugae relatively straight or slightly curved anterad; male genital cup and female basal plates exposed.................................7 7- Pronotum twice as wide as long; posterior margin of scutellum strongly concave...... Galeacius (*) - Pronotum less than twice as wide as long; posterior margin of scutellum rounded, truncate, or shallowly concave....................... Symphylus 8- Head concave dorsally, lateral and anteriormargins elevated.............................. Coptochilus - Head convex dorsally, lateral and anterior margins depressed..........................................9 9- Ostiole not attended by ruga or sulcus... Misippus - Ostiole attended by short elevated ruga... Camirus 10- Each abdominal spiracle within or subtended by callus..............................................11 - Abdominal spiracles not within or subtended by calli................................................12 11- Each abdominal spiracle located in dorsal edge of concolorous callus........................ Ascanius - Each abdominal spiracle subtended by pale callus.................................. Chelycoris 12- Distance from metacoxae to ostiole about twice the distance from ostiole to lateral margin of metapleuron.................................. Tetyra - Distance from metacoxae to ostiole subequal to or slightly greater than distance from ostiole to lateral margin of metapleuron...............13 13- Ostiolenot attendedbyrugaor sulcus........ Polytes - Ostioleattendedbyshortauriculateruga..........14 14- Abdominal sternites depressed near lateral margins................................ Pachycoris - Abdominal sternites not depressed laterally....... ................................ Orsilochides (*) Genera recorded for southern or central Brazil, probably occurring in Argentina.

Diagnosis Small to medium-sized, ovoid to elongate-ovoid in shape; most of Neotropical species are dull coloured, with a few exceptions ( Pachycoris spp., Agonosoma spp., and Augocoris spp.). Antennae 3- ( Scutellerinae ) or 5-segmented ( Pachycorinae ). Scutellum enlarged and almost entirely covering hemelytra and abdomen. Femora and tibiae unarmed. Hemelytra with frenum reduced or absent, corium and clavus membranous distally. Propleura with laminate carinae ( Schuh & Slater, 1995 ). Ostiolar peritreme of metathoracic glands absent to well developed, variable in shape, length and distance from metacoxae. External genitalia visible or concealed by seventh sternite. Aedeagus usually with three pairs of conjunctival processes. Female gonocoxites 9 (= second valvifers) fused medially ( Gapud, 1991 ), with visible fusion line ( Grazia et al. , 2008 ). Tricobothria paired. All Pachycorinae with stridulatory areas on lateral thirds of abdominal venter ( Schuh & Slater, 1995 ).

Diagnosis: The Scutellerinae are recog-nised by the following combination charac-ters: small (e.g., Figs 24b , 37 ) to very large body (e.g., Figs 19 a-d, 28a); ovoid (e.g., Figs 24b , 37 ) to elongate-ovoid ( Figs 28a, b ), sometimes strongly tapered towards terminalia; body weakly (e.g., Fig. 28a ) to heavily punctate ( Figs 24a , c-f); body brilliant, aposematically coloured, often with iridescent blue, green, orange colouration, often with contrasting markings (e.g., Figs 19 a-d, 24a, c-f, 28a-e); head subtriangular to suboval (e.g., Fig. 31a ); thoracic sterna flat (e.g., Figs 20c , 29c , 31c ); anterior margin of proepisternum weakly explanate (e.g., Figs 20c , 31c ); efferent system of metathoracic glands well-developed (e.g., Figs 20d , 25d , 31d ); peritreme mostly subreniform (e.g., Figs 20d , 39d , 42d ), sometimes large, subtriangular (e.g., Figs 25d , 29d ), or obovate (e.g., Figs 31d , 38c ); males without abdominal androconial or setose sternal glands; pregenital abdomen in both sexes without stridulatory vittae; ventral surface of pygophore caudally (e.g., Figs 32 a-h) or ventrally oriented (e.g., Figs 29e , 30e ); parameres hook-shaped (e.g., Figs 22b , 40b ); ejaculatory apparatus usually well-developed, with convoluted ventral conducting canal and oval to subelliptoid ejaculatory reservoir (e.g., Figs 30c, d , 36c, d , 40c, d ); CAI usually absent, rarely present (e.g., Figs 22c, d ); CAII usually bifurcate, with lobal sclerites (e.g., Figs 22c, d , 36c, d , 40c, d ), CAIII undivided, often U-shaped (e.g., Figs 26d , 30d , 43d ) or divided (e.g., Figs 22c, d ); spermathecal fecundation canal short or long; and, spermathecal reservoir oval.

Remarks: There is no unique character state defining the Scutellerinae , and most of the above-mentioned features are also true for the family. They are the most brilliantly coloured of all the scutellerids, but numerous Pachycorinae also exhibit colouration consistent with models of warning colouration. The male and female genitalia are also similar to taxa in other scutellerid subfamilies, such as the Elvisurinae and Sphaerocorinae, although the development of the ventral conducting canal of the ejaculatory apparatus is putatively greatest in the scutellerines, such as in Cantao parentum ( Figs 30c, d ). Many species have a U-shaped, and basally fused CAIII (e.g., Figs 26d , 30d , 43d ), but there are significant variations, particularly in Calliphara ( Fig. 22d ). Scutellerines can be separated from the Pachycorinae , and Odon-totarsinae + Tectocorinae , in lacking stridulatory vittae and androconial/setose sternal glands respectively. They can be separated from the Elvisurinae , Odontotarsinae and Sphaerocorinae superficially by their aposematic colouration (cf. dull colouration), although even in the latter subfamily some taxa exhibit moderately striking colouration, as in Sphaerocoris BURMEISTER . SCHUH & SLATER (1995) adopted LESTON’ s (1952d) definition of the Scutellerinae (including the Elvisurinae and Sphaerocorinae), but on the basis of the emarginate 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 scutellerines in the narrow sense, despite it being a ‘convenience’ group, as it helps in distinguishing the colourful Australian scutellerid species, pending a more in-depth review of the issue.

Remarks: It is beyond the scope of this work to review the Pachycorinae , which are not native to Australia; we only reviewed the introduced species, Agonosoma trilineatum in any detail. This subfamily is largely found in the Western Hemisphere, with only Deroplax MAYR and Hotea occurring in the Old World ( KIRKALDY 1909 ; SCHUH & SLATER 1995 ). KUMAR (1964) placed the Australian species, Tectocoris diophthalmus , in the Pachycorinae , on the basis of the ejaculatory apparatus; an action not followed by subsequent authors. Pachycorines are primarily diagnosed by the presence of stridulatory vittae and tibial plectra; characters that are thought to be exclusive to the Pachycorinae (e.g., SCHUH & SLATER 1995 ).

Diagnosis: The Odontotarsinae are recognised by the following combination characters: small (e.g., Figs 11 a-c) to moderately-sized species; ovoid (e.g., Figs 11 a-c), elongate-ovoid or elongate, rarely strongly tapered at terminalia; body strongly punctate; often with setigerous punctures, and sometimes with sericeous setae (Figs 13a-i), rarely with body extremely setose with elongate setae; mostly grey-brown to dark brown species (e.g., Figs 11 a-c), sometimes strami-neous, and at times with paler stripes or spots, sometimes mottled, rarely black, occasionally with patterning on scutellum (e.g., Fig. 11 a-c); larvae dull in colouration; pro- and mesothoracic sterna flat to weakly elevated, never keel-like, metasternum mostly flat, rarely a little raised; anterior margin of proepisternum weakly to moderately explanate; pronotum often subtrapezoidal, with anterolateral margins explanate, and anterior angles spinose, sometimes more rounded, with lateral margins evenly arcuate, rarely, with a medial notch, with anterior angles anteriorly projected, in front of posterior margin of eyes; external efferent system of metathoracic glands often greatly reduced, without peritreme, with prominent rounded ostiole, laterally orient-ed, evaporative areas often greatly reduced to absent, sometimes external efferent system more well developed, with short oval peritreme, elevated, bounded by moderate distribution of evaporative areas, also occurring on mesepimeron; sometimes males abdominal androconial glands present (e.g., Figs 2 a-c, 12e); pregenital abdomen without stridulatory vittae; ventral surface of pygophore caudally (e.g., Fig. 12e ) or ventrally oriented; parameres hook-shaped, often weakly (e.g., Fig. 14b ); ejaculatory apparatus weakly to moderately-developed, usually without prominent ventral conducting canal (not convoluted), ejaculatory reservoir usually small and oval (e.g., Fig. 14c ); CAI present or absent (e.g., Fig. 14c ); CAII unbranched or bifurcate (e.g., Figs 14c, d ), either sclerotized or membraneous with lobal sclerites; CAIII medially contiguous ( Fig. 11d ); spermathecal fecundation canal short; and, spermathecal reservoir oval to weakly dilated.

Remarks: STÅL (1872) described the Odontotarsinae (as Odontotarsaria) a tribe of the scutellerids, and redefined another tribe, as the Odontoscelaria, for a group of genera mostly from the Eastern Hemisphere. He separated these taxa primarily on the shape of the head (elongate versus semioval), the lateral margins of the pronotum, and degree of carination of the metathoracic sterna. SCHOUTEDEN (1904) synonymised these two taxa and the Eurygastraria sensu STÅL , and recognised the Odontotarsaria as the valid name, even though the Eurygastri-dae DALLAS and Odontoscelidae DALLAS have priority. He recognised the tribe on the basis of absence characters, such as the absence of stridulatory vittae on the pregenital abdomen, the lack of a keel-like carination on the thoracic sterna, as well as characters of general application, such as the biconvex body. KIRKALDY (1909) followed SCHOUTEDEN (1904) , and grouped the three above-mentioned suprageneric taxa within the single tribe, and included the same taxa within it. LESTON (1952d) reverted to the Stålian classification in part, by distinguishing these three groups as subtribes, within the tribe Eurygastrini . He hypothesised that these taxa were clearly differentiated from both his Scutellerini and Pachycorini on the basis of the male genitalia, suggesting that the Scutellerini-type was distinctive, and more derived from the pentatomid-type, the latter being more characteristic of pachycorines and eurygastrines. Modern Australian workers (e.g., GROSS 1975 ; MCDONALD & CASSIS 1984 ) have rejected this approach, mainly based on MC-DONALD’ s (1966) work on North American taxa, who regarded the eurygastrines as distinct from odontotarsines. SCHUH & SLATER (1995) also separated these two taxa, on the basis of the less broad scutellum (and more exposed abdominal connexiva) and reduced hindwing intervannal vein in eurygastrines. We have not evaluated either of these characters, and uncritically support the separation of the taxa. It is noteworthy that many odontotarsines and pachycorines have a reduced metathoracic peritreme, or it is absent entirely, with a distinct, laterally oriented ostiole, however these taxa are separated on the absence of stridulatory vittae in odontotarsines. The monophyly of the Odontotarsinae , however, is contentious, and we know of no character that defines them exclusively. The above diagnosis is polythetic in format and the group is retained largely as a ‘convenience’ group. There are putative infra-subfamilial clades which are worthy of examination. A group of genera related to Odon-toscelis , including those taxa with male setose sternal glands ( Table 3 ), and sometimes have rounded lateral margins of the Pronotum. This is largely a Palearctic group, but potentially includes the Australian genus Morbora (see below for discussion). Another putative clade, is represented by a group including and potentially related to Odonto-tarsus (e.g, Phimodera , Xerobia ), which lack the male sternal glands, and sometimes have excavate anterolateral margins of the Pronotum. These ideas are reminiscent of STÅL’S Odontotarsaria and Odontoscelaria, and require explicit character testing. We retain the use of the Odontotarsinae in the broad sense in this work, acknowledging the above limitations.

Remarks: STÅL (1872) first described the elvisurines as a tribal group (Elvisuraria), providing a diagnosis ( STÅL 1872 , 1873 ) based primarily on the carinate thoracic sterna. SCHOUTEDEN (1904) followed this arrangement and provided a listing of species. KIRKALDY (1909) catalogued the species and the classification of SCHOUTEDEN (1904) . The Elvisurinae are retained as a subfamily in this work, on the basis of the keel-like thoracic sterna ( Figs 4c , 7c , 10c ), which is a putative apomorphy for this taxon. In some Odontotarsinae species, the pro- and mesosterna are raised, but not to any great height, and not keel-like as in the elvisurines. Unlike, the Scutellerinae and some Pachycorinae , the adults of elvisurines are nearly all dull in colouration, aside from two species of Coleotichus ( C. blackburniae and C. bulowi ), which have a green iridescence, and some species of Chiastosternum and Solenos-thedium , which have contrasting yellow markings on the dorsum. It is noteworthy that larval elvisurines have brilliant colouration, like the larvae in the Scutellerinae . In most other characters, the elvisurines are indistinguishable from the other jewel bug subfamilies, 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 ), Pachycorinae (e.g., Camirus 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 conjunctival appendages are difficult to interpret, with the CAI absent in elvisurines, but also in many taxa in all subfamilies except the Tectocorinae . Moreover, the female terminalia 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 subfamily is restricted to the Eastern Hemisphere, with the genera distributed as follows: the monotypic genera Austrotichus and Solenotichus are endemic to Australia; Solenosthedium has seven species in the Eastern Hemisphere (from Africa to Indonesia, including Madagascar), aside from the Australian 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 Hawaii ( C. blackburniae WHITE ); the nominotypical genus Elvisura is endemic to the southern regions of the Afrotropical region, including Madagascar, to India ( 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. Vincent). He placed this taxon in the Elvisurinae , based on the conformity of the keel-like thoracic sterna with all the other members of the subfamily. We examined a single specimen of this taxon at the BMNH (label data as ‘St. Dom.’), and found that the species has elvisurine-type thoracic sterna, but possesses extensive stridulatory vittae (sublaterally on abdominal SIII-SVII), the latter being diagnostic for the Pachycorinae . On these two character systems alone, this taxon is annectant between the Elvisurinae and the Pachycorinae . Externally, this species is reminiscent of species of Coleotichus , having a rounded posterior margin of the pronotum, and densely punctate dorsum, with green iridescence (as in C. blackburniae and C. bulowi ). The external efferent system of metathoracic glands is extensive, with no peritreme evident, but with a prominent ostiole; a character state conforming to many Western Hemisphere pachycorine taxa. The external female genitalia are distinctive, with the medioposteri-or angles of the first gonocoxae narrowly excavate. There has been no examination of the male genitalia of this species. We regard this taxon as incertae sedis, pending examination of additional material, but exclude it from the Elvisurinae .

Key to Australian Scutelleridae 1 Lateral margins of thoracic sterna strongly sulcate with elevated keels ( Figs 4c , 7c , 9c ); adults dull in colouration ( Figs 3 a-e), jugal and pronotal margins entire (Figs 3a-e) ( Elvisurinae )................ 2 – Lateral margins of thoracic sterna flat, without elevated keels ( Figs 12c , 20c , 29c , 31c , 39c , 43c , 46c ); adults brightly coloured ( Figs 1 a-f, 16a, b, 19a-d, 24a,c-f, 28a-e, 45a, b), or if dull in colouration ( Figs 14 a-c), jugal and pronotal margins denticulate ( Figs 11c,d , 12 a-c), or body very small, <4 mm ( Fig. 24b , 37b )..(6) 2 Posterior margin of pronotum strongly rounded ( Fig. 3 c-e); pro- and mesosternal keels overlapping ( Fig. 7c ); peritreme of metathoracic glands elongate ( Fig. 7d ) ( Coleotichus )..................... 3 – Posterior margin of pronotum rectilinear ( Fig. 3a ), or weakly arcuate, rectilinear medially ( Fig. 3b ); pro- and mesosternal keels not overlapping ( Figs 4c , 9c ); peritreme of metathoracic glands short ( Fig. 4d ) to moderate size ( Fig. 9d )....... 5 3 Moderately sized species, males <16 mm, females <17.2 mm; labium reaching at most to metasternum; sub-anterolateral angles of scutellum without polished orange callus ( Figs 3a, b ), at most with a black spot; scutellum without patterned small black spots ( Figs 3c, d ).........4 – Large species, males> 20 mm, females> 18 mm; labium reaching abdominal SIV, with medial regions of SII-SIV correspondingly sulcate; sub-anterolateral angles of scutellum with polished orange calli ( Fig. 3e ); scutellum with pair of black spots sublateral anterior to midpoint ( Fig. 3e )..... Coleotichus excellens 4 Callosite region of pronotum with two black spots submedially ( Fig. 3d ); lateral margins of posterior 1/2 of scutellum contrastingly fuscous ( Fig. 3d ); males with abdominal setose sternal glands on SIV-SVI ( Figs 2 a-c); CAII asymmetrical ( Figs 8c, d ); CAIII medially fused ( Fig. 8d ); female terminalia co-planar; paratergites VIII large, subelliptoid... Coleotichus costatus – Callosite region of pronotum with four black spots, arranged transversely ( Fig. 3c ); lateral margins of scutellum concolorous with remainder of scutellum ( Fig. 3c ); males without abdominal setose sternal glands; CAII symmetrical; CAIII medially separated; female terminalia bi-planar, paratergites VIII angulate relative to gonocoxae I; paratergites moderately sized, subtriangular... Coleotichus artensis 5 Body elongate-oval ( Fig. 3a ); dark brown species with heavy punctation ( Fig. 3a ); ventral surface strongly setose; labium reaching posterior margin of abdominal SIII abdominal; males without abdominal sternal glands; connexival margins thickened, each segment bicoloured yellow then black ( Fig. 3a ); CAIII bifurcate, medially fused ( Fig. 4d ); spermathecal reservoir weakly dilated..... Austrotichus rugosus – Body ovoid ( Fig. 3b ); orange-brown species with shallow punctation ( Fig. 3b ); ventral surface mostly glabrous; labium reaching metasternum; males with abdominal sternal glands on SIV-SVII connexival margins greatly thickened, concolorous; CAIII not bifid, medially separated ( Figs 10c, d ); spermathecal reservoir broad, oval..... Solenotichus circuliferus 6 Both sexes with stridulatory vittae on abdominal SV-SVI ( Fig. 17e ); head large ( Figs 16a, b ); SVII enlarged, covering male ( Fig. 17d ) and female terminalia ( Pachy-corinae )........ Agonosoma trilineatum – Both sexes without stridulatory vittae on abdominal SV-SVI; head moderately sized (e.g., Figs. 19 a-d); SVII not greatly enlarged (e.g., Figs 21 a-i), terminalia of both sexes exposed, sometimes male SVIII exposed partially covering pygophore (e.g., Figs 45c , 46f ).................... 7 7 Males with androconial glands on abdominal SIV-VI ( Figs 2 d-f, g-i, 12e, 45c); external efferent system of metathoracic glands reduced (e.g., Figs 12d , 46d ); aedeagal ejaculatory apparatus without convoluted ventral conducting canal ( Figs 14c, d , 47c, d ); CAI present ( Figs 14c, d , 47c, d ); CAIII well-developed, membraneous, basally fused ( Figs 14c, d , 47c, d ).......................... 8 – Males without androconial glands, most often with abdominal sterna unadorned (e.g., Figs 21 a-i); external efferent system of metathoracic glands usually well-developed (e.g., Figs 20d , 31d ), if evaporative areas reduced, peritreme greatly enlarged, subtriangular ( Figs 25d , 29d ); aedeagal ejaculatory apparatus with convoluted ventral conducting canal (e.g., Figs 5c , 8c , 10c, d , 22c, d , 26c , 30c , 36c , 43c ); CAIII usually large, membraneous and U-shaped (e.g., Figs 26c, d , 30c, d , 40c, d , 43c, d ), rarely antler-like ( Figs 36c, d ) or digitiform ( Fig. 22c, d ) ( Scutellerinae ).... 11 8 Body small, males and females <6 mm; body dull colouration, densely punctate ( Figs 11 a-c); dorsum densely setose, with curly sericeous setae (Figs 13a, b, e, h) in-termixed with fan-like (Figs 13c, h, i) or clove-like setae (Figs 13b, f); male ab-dominal SVIII recessed, not visible ( Fig. 12e ); female gonocoxae I tripartite ( Odontotarsinae : Morbora ).......... 9 – Body large, males and female> 13 mm; body mostly orange, most often with iri-descent blue or green markings, rarely uniformly orange ( Figs 1f , 45a, b ; dorsum glabrous; male abdominal SVIII exposed, broadly covering ventral surface of py-gophore ( Figs 45c , 46e ); female gonocox-ae I undivided... Tectocoris diophthalmus 9 Anterolateral margins of pronotum and costal margins greatly expanded (Figs 11c); scutellum with broad fan-like setae (Figs 13h-i)....... Morbora schoutedeni – Anterolateral margins of pronotum and costal margins not expanded (Fig. 13a, b); scutellum with either clove-like setae (Figs 13b, 13f) or narrow fan-like setae (Fig. 13c)...................... 10 10 Dorsum with short, narrow fan-like setae, strongly tapered proximally (Fig. 13c); AI longest antennal segment; CAI branched; CAIII absent......... Morbora australis – Dorsum without fan-like setae, with clove-like setae only (Fig. 13f); AIV longest antennal segment; CAI branched; CAIII present......... Morbora hirtula 11 Body very small, <4 mm ( Figs 24b , 37 ); body with setigerous punctures ( Figs 38 a-c); head short, strongly rounded in profile ( Fig. 38b ); jugal margins rounded ( Fig. 38b ); apex of peritreme of metathoracic glands distally raised, obovate ( Fig. 38c )............... Heissiphara minuta – Body small to large species,> 8 mm; body without setigerous punctures, dorsum usu-ally glabrous, at most with simple setae; head short to elongate; jugal margins car-inate (e.g., Figs 29b , 31a, b ) or rounded (e.g., Figs 25c ); peritreme subreniform (e.g., Figs 20d , 39d , 42d ), subtriangular (e.g., Figs 25d , 29d ), rarely obovate, but if so anteriorly directed ( Fig. 31d )..... 12 12 Jugal margins straight in dorsal view ( Figs 28a , 29a ); large orange species with seven black spots on pronotum and seven spots on scutellum ( Fig. 28a ); abdominal SVIII exposed ( Figs 21e , 29e ), covering anterior margin of pygophore; CAII undi-vided ( Figs 30c, d ); female terminalia cau-dal in orientation..... Cantao parentum – Jugal margins sinuate in dorsal view (e.g., Figs 20a , 25a , 31a , 39a , 42a ); various colouring, if orange never withabove-mentioned black markings; male abdomi-nal SVIII recessed, not visible; CAII di-vided (e.g., Figs 22c, d , 26c, d , 35c, d , 40c, d , 43c, d ); female terminalia caudal or ventral in orientation............. 13 13 Dorsum densely setose ( Fig. 1d ); body iri-descent copper-dark brown ( Fig. 28c ), green ( Fig. 1f ), bicoloured dark-green/purple and yellow ( Fig. 28d ) or blue ( Fig. 28e ), often with contrasting coloura-tion (e.g., Figs 1d , 28d ); CAII(L) with perpendicular or hook-shaped apex, dis-tally denticulate ( Fig. 40c, d ) ( Lampromi-cra )........................... 14 – Dorsum usually glabrous, at most with scattered setae; CAII(L) either with digi-tiform ( Fig. 22c, d ) or subconical lobal sclerite ( Fig. 26c, d ), or bag-like ( Figs 35c, d , 36c, d ), with or without basal denticu-lation.......................... 16 14 Posterior 1/2 of scutellum yellow, re-mainder iridescent dark-green/purple ( Fig. 28d ); abdominal venter mostly iri-descent green, with yellow lateral mar-gins, inner outline of latter linear ( Fig. 21g ); CAIII medially fused post-thecal margin........................................... Lampromicra regia – Posterior 1/2 of scutellum either copper-fuscous ( Fig. 28c ) or iridescent dark green or blue ( Fig. 1f , 28e ); abdominal venter with margins of lateral colour band linear ( Fig. 21f ) or notched ( Fig. 21h ); CAIII separated post-thecal margin ( Fig. 40d ) 15 15 Dorsum uniformly copper-fuscous, some-times with iridescent green or purple sheen ( Fig. 28c ); abdominal venter most-ly concolorous with dorsum, with red lat-eral margins, inner outline of latter linear ( Fig. 21f ); anterior callus of scutellum punctate, always fuscous ( Fig. 28c ); CAII(L) apex hook-like ( Figs 40c, d ).................. Lampromicra aerea – Dorsum mostly iridescent green ( Fig. 28e ) or blue, often with anterior orange callus on scutellum ( Fig. 1d ); abdominal venter mostly iridescent dark green or blue, with orange lateral margins, inner outline of latter notched ( Fig. 21h ); anterior callus of scutellum impunctate ( Figs 1f , 28e ); CAII(L) apex linear.............................. Lampromicra senator 16 AII(b) minute, shortest antennal seg-ment; AII(a) arcuate, more so in males; dorsum strongly iridescent green with or-ange and fuscous markings ( Fig. 28b ); lat-eral areas of mesosternum striate; crown of paramere weakly curved, apex truncate ( Fig. 43b ); CAII(L) with an elongate ar-cuate lobal sclerite ( Figs 43c, d )..................... Scutiphora pedicellata – AI or AII(a) shortest segment; AII(a) lin-ear; body usually red or orange with con-trasting markings, if iridescent green without orange markings; lateral areas of mesosternum smooth; crown of paramere strongly hooked (e.g., Fig. 36b ); CAII ei-ther undivided, or with base membrane-ous post-thecal margin (e.g., Figs 35c, d , 36c, d ), if entirely sclerotized, oval in shape.......................... 17 17 Body small to moderate size, males <12.5 mm, females <13 mm; dorsum densely punctate ( Figs 24 c-f); jugal margins cari-nate ( Figs 31a, b ); hemelytral membrane tip not exposed beyond scutellum (Figs 24c-f); peritreme of metathoracic glands obovate, elongate, laterally recurved to-wards head ( Fig. 31d ); ventral surface of male pygophore caudally oriented (Figs 31e, 32b, f, h) ( Choerocoris )........ 18 – Body moderate size to large, males> 13 mm, females> 13.5 mm; dorsum punctate to impunctate; hemelytral membrane tip exposed beyond scutellum ( Figs 19 a-d); peritreme of metathoracic glands either subtriangular ( Fig. 25e ) or subreniform ( Fig. 20d ); ventral surface of male py-gophore ventrally oriented ( Figs 20 a-c) or caudally oriented.............. 21 18 Callosite region of pronotum medially impunctate ( Fig. 24d ); callosite region de-marcated posteriorly by transverse furrow ( Fig. 24d ); scutellum red with pair of sub-medial, iridescent blue, subtriangular markings ( Fig. 24d ), anterior margin with iridescent blue marking; apex of scutel-lum without markings ( Fig. 24d ); abdom-inal venter mostly red with lateral and submedial dark blue markings ( Figs 32e, f ); CAII(M) and CAII(L) roughly sym-metrical.......... Choerocoris paganus – Pronotum densely and uniformly punc-tate ( Figs 24c, e, f ); callosite and disc re-gions of pronotum co-planar ( Figs 24c, e, f ); scutellum orange or red with three prominent iridescent blue regions, always with V-shaped distal markings ( Figs 24c, e, f ); abdominal venter mostly orange or red, with ( Figs 32a, b ) or without subme-dial darker markings ( Figs 32c, d, g, h ); CAII(M) and CAII(L) significantly dif-ferentiated ( Figs 35c, d , 36c, d )..... 19 19 Jugae orange ( Fig 24c ); abdominal venter mostly orange with lateral blue or green markings ( Figs 32c, d ); apex of CAII(M) with weak sclerotization ( Fig. 35c, d ); CAIII contiguous medially, lightly sclero-tized, lobe-like ( Figs 35c, d ); vesica with-out distal process beyond secondary gono-pore ( Fig. 35c, d )............................. Choerocoris lattini nov.sp. – Head mostly iridescent dark green or blue ( Figs 24e, f ); abdominal venter either yel-low or orange, with ( Fig. 32a, b ) or with-out submedial dark ( Figs 32g, h ) blue markings; apex of CAIII with prominent triangular lobal sclerite ( Figs 36c, d ); CAIII separated, as antler-like and heavi-ly sclerotized processes ( Figs 36c, d ); vesi-ca with subdistal paired process ( Figs 36c, d ) ............................ 20 20 Pregenital abdominal venter mostly yel-low with submedial and lateral black bands ( Figs 32a,b ); posterior angles of ab-dominal connexiva V-VII nodulate, yel-low ( Figs 32a, b ); female paratergites IX greatly expanded, much larger than gono-coxae I ( Figs 33e, f ); gonocoxae I greatly reduced, posterior margin deeply excavate ( Figs 33e, f )... Choerocoris grossi nov.sp. – Pregenital abdominal venter mostly yel-low with lateral black band only (Figs 32g, h); posterior angles of abdominal connexiva V-VII weakly swollen at most, fuscous ( Figs 32g, h ); female paratergites IX not greatly enlarged, smaller than gonocoxae I ( Figs 33c, d ); gonocoxae I well-developed, posterior margin weakly excavate ( Figs 33c, d )............................ Choerocoris variegatus 21 Hemelytral membrane barely surpassing tip of scutellum ( Fig. 24a ); dorsum mostly fuscous, with orange/red and iridescent dark green markings ( Figs 24a ); AII(a) moderately short, AII(b) at most 2.5x> AII(a); peritreme of metathoracic glands greatly expanded laterally, subtriangular, 1-1.5x longer than wide ( Figs 25d ); evap-orative areas restricted, not extending to mesepimeron ( Fig. 25d ); CAI absent (Figs 26c); female terminalia caudal in orienta-tion.................. Calliscyta stalii – Hemelytral membrane clearly surpassing tip of scutellum ( Figs. 19 a-d); dorsum mostly iridescent green, orange, or orange and blue, with various markings (Figs 19a-d); AII(a) minute, AII(b) more than 6x> AII(a) length; peritreme of metatho-racic glands subreniform, tapered distally, 3x or more longer than wide ( Fig. 20d ); evaporative areas extensive, extending to mesepimeron ( Fig. 20d ); CAI present ( Figs 22c, d ); female terminalia ventral in orientation ( Calliphara )........... 22 22 Eye length 1/2 head length ( Fig. 19d ); head orange-red to copper with fuscous markings ( Fig. 19d ); pronotum iridescent copper, with calli most often fuscous ( Fig. 19d ); scutellum iridescent copper, with fuscous spot at tip ( Fig. 19d ); exocorium iridescent green ( Fig. 19d ); posterior an-gles of abdominal SVI-SVII spinose; CAI bifurcate, with CAI(D) elongate, digiti-form ( Fig. 22c, d ); CAIII divided (Figs 22c, d); vesica with apical hood ( Fig. 22c, d )................. Calliphara regalis – Eye length <1/2 head length ( Figs 19 a-c); head concolorous, iridescent dark green or blue; pronotum orange or irides-cent green, rarely with calli fuscous; scutellum broadly bicoloured ( Fig. 19a ), or orange with narrow distal iridescent green marking ( Fig. 19b ); posterior an-gles of abdominal sterna without spines ( Fig. 21c ), or SIV-SVI ( Fig. 21b ) or SIV-SVII spinose ( Fig. 21a ); CAI and CAIII divided or undivided; vesica with or without processes................ 23 23 Scutellum either red and iridescent dark blue ( Fig. 19a ) or mostly orange ( Fig. 19b ); legs concolorous, fuscous; abdominal venter impunctate ( Figs 21a, b ), sometimes rugose laterally; spinose posterolateral angles of abdominal sterna SIV-SVI ( Fig. 21b ) or SIV-SVII ( Fig. 21a ).... 24 – Scutellum mostly iridescent green, each with seven black spots ( Fig. 19c ); femora mostly orange, apex narrowly fuscous; tibiae and tarsi fuscous; abdominal venter punctate ( Fig. 21c ); posterior angles of abdominal sterna not spinose ( Fig. 19c ).................. Calliphara nobilis 24 Scutellum mostly orange, with extreme caudal tip iridescent green ( Figs 1a , 19b ); dorsum with moderately deep punctation; abdominal pregenital venter mostly iridescent green, laterally red to orange-red, rugose ( Fig. 21b ); posterolateral angles of abdominal SIV-SVI spinose ( Fig. 21b )................ Calliphara imperialis – Scutellum broadly bicoloured, anterior 1/2 orange to orange-red, remainder iridescent dark blue, with w-shaped anterior outline ( Fig. 19a ); dorsum with shallow punctation; abdominal pregenital venter orange, laterally smooth ( Fig. 21a ); posterolateral angles of abdominal SIV-SVII spinose ( Fig. 21a )........................... Calliphara dimidiata cruenta

Description: Body elongate to elongate-ovoid; moderately to strongly biconvex, sometimes dorsum more rounded than venter; mostly large species, between 8-20 mm, smallest species <4 mm, largest species> 20 mm; body either dull brown to black in colouration (e.g., Figs 3 a-e), or brightly coloured, with iridescent hue and often with contrasting markings (e.g., Figs 19 a-d, 24a-f, 28a-e); body often punctate, with shallow (e.g., Figs 7a ), to deep punctures (e.g., Figs 4a , 12a , 31a , 37 ), usually regularly distributed, sometimes punctures coalesced (e.g., Fig. 3a ); body less commonly rugo-punctate on lateral aspects of pronotum and abdominal venter; body mostly with sparse distribution of simple setae, rarely with dense distribution of simple setae (e.g., Fig. 1d ), rarely with setigerous ( Figs 31a , 38a ) or scale-like setae (e.g., Figs 13c, h, i). Head: usually triangular to subtriangular (e.g., Figs 4a , 7a , 17a ) sometimes suboval (e.g., Fig. 9a ); jugae large, not exceeding the anterior edge of clypeus (e.g., Figs, 4a, 20a, 42a, 46a); lateral margins of jugae most often excavate (e.g., Figs 20a , 25a ), sometimes linear (e.g., Figs 29a , 46a ), rounded (e.g., Figs 7b , 25c , 38b , 39b , 42b ) or carinate (e.g., Figs 29b , 31a, b ) in profile; clypeus well-developed, lateral margins reaching between frons (e.g., Figs 4a , 7a , 9a , 31a ) and anterior aspect of vertex (e.g., Figs 25a , 38a , 42a ); vertex weakly convex; lorae triangular, not visible from above, with strongly demarcated margins; bucculae large, narrow, subparallel, usually reaching near base of head (e.g., Figs 4c , 7c , 20c , 31c , 46c ); gula short, convex. Eyes: contiguous with pronotum (e.g., Figs 3 , 24 , 45a, b ); usually moderately sized, ≥ 1/3 of head length, sometimes large, about 1/2 head length ( Fig. 19d ); pair of ocelli, removed from posterior margin of head (e.g., Figs 3 , 24 , 45 ). Antennae: inserted anteriad of ventral margin of eyes (e.g., Figs 7b , 9b , 20b , 29b , 31b , 38b , 39b , 46b ); four-segmented, AII divided, relatively short; segments linear, cylindrical, AII(b)-AIV often flattened; AII rarely arcuate; AI short, sometimes AII(a) and AII(b) shortest segment; AIV often longest segment. Labium: usually reaching between apices of mesocoxae to abdominal SIV, rarely longer; LI reaching posterior margin of head; LII usually longest segment; LII sometimes laterally bicompressed; LIII-LIV sometimes dorsoventrally flattened. Pronotum: large, trapeziform; anterior margin weakly (e.g., Figs 3a , 7a ) to strongly concave (e.g., Fig. 9a ); anterolateral margins elongate, weakly (e.g., Figs 16a , 25b ) to strongly divergent (e.g., Figs 9a , 19 a-d, 45a,b, 46a) posteriorly, anterolateral margins usually linear (e.g., Figs 3 a-e, 19a-d), rarely weakly convex (e.g., Figs 24c, d, e ) or concave (e.g., Figs 28b , 45a, b ), carinate (e.g., Fig. 25d ) or rounded in profile; posterolateral margins short, most often weakly convex; posterior margin usually rectilinear (e.g., Figs 3a , 16a, b , 19 a-d, 24a-f), rarely rounded (e.g., Figs 3 c-e), if so covering anterior margin of scutellum, rarely weakly excavate (e.g., Fig. 28a ); callosite region with sublateral triangular markings, most often not strongly demarcated, co-planar with pronotal disc (e.g., Figs 3 a-e), rarely with transverse furrow (e.g., Figs 28 c-e). Scutellum: greatly enlarged, shield-like, U-shaped, covering most of hemelytra and abdomen (e.g., Figs 3 a-e, 16a, b, 19a-d, 24a-f, 28a-e, 45a, b), exocorium always partly exposed, base of clavus sometimes exposed, tip of hemelytral membrane sometimes visible beyond scutellum at rest (e.g., Figs 19 a-d, 28a); strongly convex, often strongly declivent posteriorly, beyond midpoint; anterior margin sometimes with sublateral foveae (e.g., Figs 24a , 28a ); lateral margins weakly to strongly rounded, sometimes strongly divergent posteriorly; tip rounded, rarely truncate ( Fig. 24a ). Frena: greatly reduced to absent. Hemelytra: membrane with ten (e.g. Fig. 24c ) or more longitudinal veins. Thoracic pleura: anterior margin of proepisternum weakly (e.g., Figs 4c , 7c , 20c , 31c ) to strongly (e.g., Figs 7c ) explanate, sometimes excavate at eye (e.g., Fig. 7c ), sometimes entire and linear ( Figs 17b , 39c , 46 b-c); mesepimeron subrectangulate, most often with evaporative areas on posterior margin, bounding metathoracic spiracle (e.g., Figs 4d , 7d , 9d , 12d , 17c , 20d , 31d , 38c , 42d ), sometimes extending to meso-supracoxal lobe, sometimes absent (e.g., Figs 25d , 46d ), posterior margin sometimes swollen (e.g., Fig. 12d ); metepisternum usually with well-developed external efferent system of metathoracic glands (e.g., Figs 4d , 7d , 9d , 17d ,) usually covering at least 1/2 of segment, rarely reduced (e.g., Figs 12d , 42d , 46d ); ostiole small (e.g., Fig. 4d , 46d ) to moderately large, oriented near anterior margin of segment; peritreme very small (e.g., Fig. 4d ) to greatly enlarged (e.g., Fig. 7d ), either linear (e.g., Figs 7d ), linear with apex anteriorly recurved (e.g., Figs 17c , 31d ), subreniform (e.g., Figs 20d , 39d , 42d ), obovate ( Fig. 38c ) or greatly enlarged and subtriangular ( Figs 25d , 29d ), peritreme often medially sulcate (e.g, Figs 7d ); evaporative areas reduced (e.g., Figs 25d , 29d , 42d , 46d ) to extensive (e.g, Figs 4d , 7d , 9d , 17c , 31d ), often extending laterally beyond tip of peritreme. Thoracic sterna: usually flat (e.g., Figs 12c , 20c , 29c , 31c , 39c , 42c , 46c ), sometimes lateral margins strongly raised, keel-like (e.g., Figs 4c , 7c , 9c ), rarely with pro- and mesosternal keels overlapping ( Fig. 7c ). Legs: femora fusiform; tibiae terete, tarsi three-segmented. Pregenital Abdominal Venter: SII-SVII always visible ( Figs. 21 a-d, f-i), SVIII most often recessed, rarely exposed in males ( Figs 21e , 45c ), covering posterior margin of pygophore ( Figs 29e , 46e ); sometimes posterolateral angles with acuminate spine (e.g., Figs 21a, b ); males with sternal glands sublaterally, ranging between SIII-SVII ( Figs 2 a-i, 12e); sometimes males and females with stridulatory region on SV-SVI ( Figs 17 e-f). Male Genitalia: pygophore moderate size to large, ventral surface caudally (e.g., Figs 4e , 9e , 12e , 31e ) or ventrally oriented (e.g., Figs 20e , 29e , 42e , 46e ); ventral margin convex, emarginate or sublinear; genital opening moderately broad to broad, with setose regions usually on dorsal margin (e.g., Figs 22a , 43a ), and often on ventral (e.g., Figs 22a , 46f ) and lateral margins (e.g., Figs 4f , 7f , 8a , 31f , 43a , 46f ); parameres moderate size (e.g., Figs 14b , 40b , 43b ) to large (e.g., Figs 36b , 47b ), symmetrical, with columnar stem, and weakly (e.g., Figs 14b , 26b , 35b , 40b , 43b ) to strongly hooked (e.g., Figs 8b , 10b , 18b , 22b , 47b ) crown, usually with apex of stem setose, and sometimes with flange (e.g., Figs 5b , 8b , 35b , 36b ); aedeagus with mesial U-shaped phallobase, with round capitate processes; aedeagus box-like, often heavily sclerotized; phallotheca cylindrical (e.g., Figs 5c , 26c ) to subconical (e.g., Figs 8c , 10c , 22c ), thick, sometimes with pair of subdistal thorn-like processes ( Fig. 40c ), or areas of sclerotization (e.g., Figs 5c, d , 30c , 36c ); ductus seminis proximalis narrow, membraneous, without ribbing (e.g., Figs 5c , 8c , 10c, d , 14c , 22c, d , 26c , 30c , 36c , 43c ) sometimes thick-er ( Fig. 18c ), rarely bounded by sclerotized process of vesica ( Fig. 30c ) or ligamentary tubule (e.g., Figs 18c , 47c, d ); ejaculatory apparatus elaborate (e.g., Figs 5c , 8c , 10c, d , 14c , 22c, d , 26c , 30c , 36c , 43c ), moderately to heavily sclerotized, most often with convoluted ventral conducting canal (e.g., Figs 5c , 8c , 10c, d , 22c, d , 26c , 30c , 36c , 43c ), with 2-12 convolutions; ejaculatory reservoir small (e.g., Fig. 22c ), moderate size (e.g., Figs 5c , 8c , 10c , 14c , 43c ) to large (e.g., Figs 18c , 22c, d , 26c , 30c , 35c , 36c , 40c ), elliptoid to oval, sometimes greatly reduced and heavily sclerotized ( Fig. 47c ); dorsal conducting canal simple, narrow to relatively broad (e.g., Figs 5c , 8c , 10c, d , 22c, d , 26c , 30c , 36c , 43c ); ductus seminis distalis usually narrow, rarely expanded; vesica rarely membraneous ( Fig. 18c, d ), usually sclerotized, moderate size (e.g., Figs 5c, d , 22c, d , 36c, d , 47c, d ), or elongate (e.g., Figs 8c, d , 10c, d , 26c, d , 30c, d , 35c, d , 40c, d , 43c, d ), arcuate (e.g., Figs 10c, d , 14c, d , 43c, d , 47c, d ), sinuate (e.g., Fig. 5c, d , 8c, d , 30c, d , 35c, d , 40c, d ), rarely subdistally strongly incrassate (e.g., Figs 30c , 35c ), sometimes with subdistal processes (Fig, 36c, d), or with distal process above secondary gonopore ( Figs 22c, d ); CAI most often absent (e.g., Figs 5c, d , 8c, d , 10c, d , 26c, d , 30c, d , 35 , d, 36c, d , 40c, d , 43 , d), if present medially fused at base ( Fig. 14c, d , 47c, d ), or bifurcate with ventral and dorsal branches (e.g., Figs 22c, d ); CAII sometimes undivided (e.g., Figs 36c, d , 47c, d ), usually bifurcate (e.g., Figs 5c, d , 8c, d , 10c, d , 26c, d , 30c, d , 35 , d, 36c, d , 40c, d , 43c, d ), CAII(L) often basally membraneous, rarely without lobal sclerites (e.g., Fig. 35c, d ), usually with apical lobal sclerite, either short, subconical (e.g., Figs 10c, d , 26c, d ), hooked with spicules (e.g., Fig. 40c, d ), or digitiform (e.g., Figs 8c, d , 43c, d ), sometimes entirely sclerotized (e.g., Figs 22c, d ), sometimes spiculate, CAII(M) usually membraneous with apical lobal sclerite, either short (e.g., Figs 5c, d ), triangular (e.g., Figs 36c, d ), digitiform lobal sclerite (e.g., Figs 26c, d ), hooked (e.g., Figs 40c, d ), or with bifid sclerotized process (e.g., Figs 22c, d , 43c, d ); CAIII either separated, heavily sclerotized, moderate size, arcuate (e.g. Figs 10c, d ), elongate and sclerotized (e.g., Figs 22c, d ), medially fused, small (e.g., Figs 8c, d ) or large and U-shaped (e.g., Figs 26c, d , 30c, d , 40c, d , 43c, d ), or antler-like ( Figs 36c, d ), or medially fused and membraneous (e.g., Figs 14c, d , 47c, d ), sometimes bifurcate, with dorsal and ventral branches (e.g., Figs 5c, d ), rarely absent ( Fig. 18c ). Female Terminalia: either co-planar or bi-planar (e.g., Figs 33d, f ); either ventrally or caudally oriented, sometimes recessed; paratergites VIII moderate size to large, subtriangular to subelliptoid, usually strongly tapered towards midline; paratergites IX small (e.g., Fig. 33b ) to large (e.g., Figs 33e, f ), usually ovoid and broader in middle, rarely tapered medially; gonocoxae I most often entire, rarely subdivided, usually moderate size, greater than paratergites IX, subtriangular, posterior margin rectilinear to weakly excavate, sometimes depressed medially, and often with medial margins raised; first and second gonapophyses membraneous; first and second rami usually developed. Spermatheca: proximal fecundation canal short to elongate, membraneous, sometimes strongly sclerotized basally; reservoir usually broad, oval, often ribbed, rarely weakly dilated, membraneous; distal fecundation canal usually short, rarely elongate, sometimes heavily sclerotized, usually with proximal and distal flanges, bulb small, oval, sometimes bilobed. Larvae: dull or iridescent colouration; dorsal abdominal gland openings between terga III-IV, IV-V and V-VI.

Diagnosis: The Tectocorinae are recognised by the following combination of characters: usually large species ( Figs 45a, b ); aposematic colouration; elongate-ovoid ( Figs 1f , 45a, b ), shallowly punctate; elongate head ( Fig. 46a ); thoracic sterna flat ( Fig. 46c ); anterior margin of proepisternum linear ( Fig. 46c ); efferent system of metathoracic glands vestigial, with rectilinear peritreme ( Fig. 46d ); males with abdominal androconial glands on SIV-SVI ( Figs 2 g-i, 45c); pregenital abdomen in both sexes without stridulatory vittae; ventral surface of pygophore ventrally oriented (e.g., Figs 46c ); parameres large, strongly hook-shaped ( Fig. 47b ); ductus seminis proximalis broad, bounded by ligamentary tubule ( Figs 47c, d ); ejaculatory apparatus box-like, heavily sclerotised; without convoluted ventral conducting canal ( Figs 47c, d ); three conjunctival appendages present, mostly membraneous ( Figs 47c, d ); CAII undivided with conical lobal sclerite ( Figs 47c, d ); females lacking interlocking rami; spermathe-cal fecundation canal short; spermathecal reservoir membraneous, oval; spermathecal pump minute, without flanges.

1 Antennae with three segments, abdomen without stridulating areas................ ( Scutellerinae ) Augocoris Burmeister - Antennae with five segments, abdomen with stridulating areas....................................Pachychorinae (2) 2 Semirectangular or semiquadrangular head, dorsally flattened, with concave mandibular plates and elevated anterior and lateral margins................................................................... Coptochilus Amyot and Serville - Head variable in shape, dorsally semi-flattened, convex or slightly convex with anterior and lateral margins not elevated... 3 3 Distance from metacoxa to ostiole shorter than the distance from ostiole to the lateral margin of metapleura............. 4 - Distance from the metacoxa to the ostiole greater than or nearly equal to the distance from the ostiole to the lateral margin of the metapleura....................................................................................... 6 4 Pronotum with a median transverse impression, usually conspicuous; head declivent; punctuation thick and dense, usually with short and decumbent setae.............................................................................. 5 - Pronotum without transverse impression; head generally straight; punctuation variable, rarely with setae and, if present, then generally long and erect................................................................................ 9 5 Head strongly declivent; bucculae with distinct lobe anteriorly, lobe strongly truncated posteriorly; coloration black; stridulatory areas not extending to the seventh sternite................................................ Acantholomidea Sailer - Head slightly to moderately declivent; bucculae gently rounded, not posteriorly truncated; coloration brown, usually mottled; stridulating areas often extending to the seventh sternite............................................. Camirus Stål 6 Dorsum covered with long, thin setae; head shorter than pronotum; antennomeres 4 and 5 distinctly flattened.... Dystus Stål - Dorsum generally glabrous or at most with short and decumbent setae; other characters variable....................... 7 7 Small-sized species (≤ 12 mm ); short or underdeveloped ostiolar peritreme...................... Orsilochides Kirkaldy - Large species (> 12 mm ); no ostiolar peritreme, or if present, then well developed.................................. 8 8 Anterolateral region of head truncated and oblique; scutellum not as wide as abdomen; distance from ostiole to metacoxa about 1.5 times the distance from ostiole to the lateral margin of metapleura; if less, femora are provided with a black ring or a ring of black maculae......................................................................... Tetyra Fabricius - Anterior region of head rounded; scutellum equal or nearly equal in width to the abdomen; distance from ostiole to metacoxa nearly equal to or slightly greater than the distance from ostiole to the lateral margin of metapleura; femora uniform in color.................................................................................. Pachycoris Burmeister 9 Ostiolar peritreme occupying less than one-third of the width of the metapleuron or length of the ostiolar peritreme less than half the distance from the ostiole to the lateral margin of the metapleura................................ Tiridates Stal - Ostiolar peritreme occupying one-third or more of the width of the metapleuron or length of the ostiolar peritreme greater than half the distance from the ostiole to the lateral margin of the metapleura......................................... 10 10 Ostiolar peritreme oblique, with the curved apical third gradually expanded and forming an almost right angle................................................................................................. Sphyrocoris Mayr - Ostiolar peritreme deeply or slightly concave, with the apical third curved or slightly curved but in both cases not expanded. .................................................................................................. 11 11 Ostiolar peritreme cleft and distinctly curved, with raised sides; posterolateral angles of the last abdominal sternites not developed into small spines............................................................... Homaemus Dallas - Ostiolar peritreme curved or slightly curved, but with its sides not elevated; usually with posterolateral angles of at least the last three abdominal sternites developed into small spines........................................................ 12 12 Elongated ostiolar peritreme almost reaching the anterior margin of the metapleura, with its sides parallel or tapering slightly toward the apex........................................................................ Brailovskylus Eger - Elongated ostiolar peritreme but clearly not reaching the anterior margin of the metapleura.......................... 13 13 Axe-shaped or hammer-shaped parameres.................................................... Symphylus Dallas - Hook-shaped parameres................................................................ Stethaulax Bergroth