Scutelleridae, Leach, 1815

Gerry Cassis & Loren Vanags, 2006, Jewel Bugs of Australia (Insecta, Heteroptera, Scutelleridae), Denisia 19, pp. 275-398 : 293-300

Family Scutelleridae

Scutellerida LEACH 1815: 123 (suprageneric group) Scutelleroides: DALLAS 1851: 3 (superfamily; infrafamilial classification) Pentatomidae , Scutellerina: STÅL 1872: 32 (classification); STÅL 1873: 3 (classification) Pentatomidae , Scutelleridae (sic): LETHIERRY & SEVERIN 1893: 15 (catalogue) Pentatomidae , Scutellerinae: SCHOUTEDEN 1904: 1 (monograph); KIRKALDY 1909: 263 (catalogue); LESTON 1952d: 13 (subfamily) Scutelleridae : MCDONALD 1966: 67, 68 (genitalia); MCDONALD & CASSIS 1984: 550 (revision); CASSIS & GROSS 2002: 580 (catalogue)

Diagnosis: Scutellerids are recognised by the following combination of characters: body moderately to strongly biconvex (e.g., Fig. 1 ); apices of jugae and anterior edge of clypeus subequal in length (e.g., Figs 4a , 7a , 20a , 46a ); bucculae flattened, sub-parallel (e.g., Figs 7c , 25b , 46c ); antennae four-segmented, AII divided, distal segments often flattened; labium four-segmented, most often reaching between metacoxae and abdominal SIV; pronotum trapeziform, with anterolateral and posterolateral margins (e.g., Figs 3 , 16 , 19 , 45a, b ), or rounded lateral margins ( Figs 16 a-c); callosite region weakly demarcated; scutellum enlarged, covering most of abdomen and wings, with exocorium and clavus minimally exposed proximally (e.g., Figs 3 , 16 , 19 , 45a, b ); forewing membrane sometimes exposed beyond tip of abdomen (e.g., Figs 19 a-d); frena reduced or absent; hemelytral membrane with 10 or more longitudinal veins; external efferent system of metathoracic glands present (e.g., Figs 4d , 17c , 20d , 46d ); ostiole placed anteriorly on metepisternum; evaporative areas present on metepisternum, and most often on mesepimeron (e.g., Figs 4d , 12d , 20d , 31d ); parameres with a columnar stem, and usually with a hook-shaped crown (e.g., Figs 5b , 22b , 47b ); aedeagus most often with an ejaculatory apparatus (e.g., Figs 5c, d , 26c, d , 30c, d ); first and second gonapophyses membraneous; spermathecal reservoir without a sclerotized rod; and, spermathecal bulb round without processes.

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.

Diversity and distribution: The Scutel-leridae are composed of about 80 genera and 500 species ( LATTIN 1964; Zoological Record 1965-2005). Of the scutellerid subfamilies, the nominotypical Scutellerinae are the most diverse and are primarily found in the Old World tropics, with only one genus, Au-gocoris BURMEISTER , known from the Western Hemisphere. The Pachycorinae are also diverse, with over 100 described species, including numerous undescribed taxa, and are primarily found in the Western Hemisphere, mostly in the Neotropical zoogeographic region, with a few genera also represented in the Afrotropical ( Deroplax MAYR ) or Afrotropical and Oriental regions ( Hotea AMYOT & SERVILLE ). The Odontotarsinae are the next most diverse suprageneric group, and have putatively the broadest distribution of all the scutellerid subfamilies, with a near cosmopolitan distribution, although more than 80 of the species of this subfamily are found in the Palaearctic region. The Elvisurinae and the Sphaerocorinae are small subfamilies, with less than 25 species, which are found primarily in the Old World tropics. The Eurygastrinae and Tecto-corinae are currently defined as monogeneric subfamilies, of the Eastern Hemisphere.

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

Figures

Fig. 1: Examples of Australian Scutelleridae species and their host plant species (a) adult aggregation of Calliphara regalis (b) Cantao parentum on Mallotus philippensis (c) Choerocoris paganus on Dodonaea viscoca (d) Lampromicra senator on unidentified host plant (e) Scutiphora pedicellata on unidentified host plant (f) Tectocoris diophthalmus on Brachychiton acerifolius.

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 = mesepimeron; Met = metepisternum; Ms(k) = mesosternal keel; Pe = peritreme; Pe(k) = proepisternal keel, ventral view; P(r) = right paramere; Pr = proctiger. Scale bars = 1 mm.

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.

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.

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 .

Fig. 25: Scanning electron micrographs of key characters of Calliscyta stalii (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.

Fig. 3: Habitus of Australian Elvisurinae species (a) Austrotichus rugosus, Oi (b) Solenotichus circuliferus, Oi (c) Coleotichus artensis, Oi (d) Coleotichus costatus, Oi (e) Coleotichus excellens, Oi. Scale bars: 1 mm.

Fig. 16: Habitus of introduced Pachycorinae species (a-b) Agonosoma trilineatum (a) ♂ (b) ♀. Scale bar = 1 mm.

Fig. 19: Habitus of Australian Calliphara species (a) Calliphara dimidiata cruenta, Oi (b) Calliphara imperialis, Oi (c) Calliphara nobilis, Oi (d) Calliphara regalis, Oi. Scale bar = 1 mm.

Fig. 45: Habitus and abdominal venter of Tectocoris diophthalmus (a) ♂ (b) ♀ (c) abdominal venter.

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.

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.

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.

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.

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.

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.

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.

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.

Fig. 37: Habitus of holotype of Heissiphara minuta nov.sp., ♀.

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.

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.

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.

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.

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.

Fig. 24: Habitus of Australian Scutellerinae species (a) Calliscyta stalii, ♀ (b) Heissiphara minuta nov.sp., ♀ (c) Choerocoris lattini nov.sp., ♀ (d) Choerocoris paganus, ♂ (e) Choerocoris grossi nov.sp., ♂ (f) Choerocoris variegatus, ♂. Scale bar = 1 mm.

Fig. 28: Habitus of Australian Scutellerinae (a) Cantao parentum, Oi (b) Scutiphora pedicellata, ♀ (c) Lampromicra aerea, Oi (d) Lampromicra regia, Oi (e) Lampromicra senator, Oi. Scale bars = 1mm.

Fig. 21: Colouration of Abdominal Venter in Australian Scutellerinae (a) Calliphara dimidiata cruenta (b) Calliphara imperialis (c) Calliphara nobilis (d) Calliscyta stalii (e) Cantao parentum (f) Lampromicra aerea (g) Lampromicra regia (h) Lampromicra senator (i) Scutiphora pedicellata.

Fig. 2: Abdominal Sternal Glands of Scutelleridae (a-c) Coleotichus costatus (a) abdominal venter, sternal glands on SIV-SVI (b-c) higher magnification of sternal glands (d-f) Morbora schoutedeni (d) androconial glands on SIV-SVI (e) higher magnification of microsculpture medial to androconial glands (f) higher magnification of apex cuticular duct of glandular unit of androconial glands (g-i) Tectocoris diophthalmus (g) androconial glands on SIV-SVI (h) higher magnification of androconial glands on abdominal SVI (i) higher magnification of apex of cuticular ducts of glandular units of androconial glands. Abbreviations: Ag = Androconial glands; Cd = cuticular ducts; Sg = sternal glands. Scale bars = 0.1 mm.

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.

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.

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.

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.

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.

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.

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.

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.

Fig. 33: Female Terminalia of Choerocoris species (a) Choerocoris lattini nov.sp., ventral view (b) Choerocoris paganus, ventral view (c-d) Choerocoris variegatus (c) ventral view (d) lateral view (e-f) Choerocoris grossi nov.sp. (e) ventral view (f) lateral view. Abbreviations: PVIII = eighth paratergites; PIX = ninth paratergites; GXI = gonocoxae I.

Fig. 11: Habitus of Morbora species (a) M. australis (b) M. hirtula (c) M. schoutedeni. Scale bars = 1 mm.

Fig. 32: Colouration of Abdominal Venter of Choerocoris species (a-b) Choerocoris grossi 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 variegatus (g) ventral view (h) lateral view.