Heraeus
Creators: name: Pablo M. Dellapé
name: María C. Melo
name: Thomas J. Henry
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hemiptera
Rhyparochromidae
Heraeus

description: KEY TO THE SPECIES OF HERAEUS
1. Hemelytra nearly uniformly coloured, without a large pale spot at apex of corium (or with a small, pale subapical corial spot, with apex darker; Fig. 3 A–E); head smooth (coquilletti group) ................................................... 2
– Hemelytra more heterogeneously coloured, often with a large pale spot at apex of corium (Fig. 3 F, K); head rugose. ......................................................................................................................................................6
2. Pronotum and scutellum with long erect setae ....................................................................................... 3
– Pronotum and scutellum without long erect setae ................................................................................... 4
3. Distiflagellomere pale brown, with a pale band; hemelytra with only short, recumbent setae (Mexico; Fig. 3 D) ... ..................................................................................................................... Heraeus itzelae sp. nov.
– Distiflagellomere yellowish brown, without a pale band; hemelytra with long, erect setae (Honduras and Mexico; Fig. 3 E).......................................................................................................... Heraeus setosus sp. nov.
4. Head, pronotum, and hemelytra shiny (Mexico and USA; Fig. 3 B).................... Heraeus coquilletti Barber, 1914
– Head shiny, contrasting with a dull pronotum and hemelytra ................................................................... 5
5. General colouration dark brown; distiflagellomere mostly white, with the apex dark; profemur mostly dark brown (Costa Rica, Honduras, and Nicaragua; Fig. 3 C) ................................................... Heraeus costalis sp. nov.
– General colouration cinnamon brown; distiflagellomere uniformly brown; profemur pale yellowish brown (Mexico and USA; Fig. 3 A)............................................................................... Heraeus cinnamomeus Barber, 1948
6. Hemelytron with a distinct, white, subapical spot on corium and a white mark on apex of membrane; species usually glabrous (guttatus and caliginosus groups; Fig. 3 F–K)..............................................................................7
– Hemelytron with different combination of spots; if subapical corial spot present, shape is irregular and the colour pattern is more complex; species often setose (Figs 15, 25, 36).................................................................12
7. Labium extending beyond metacoxae; posterior pronotal lobe with four small pale spots strongly contrasting with dark background (Dominican Republic and Haiti; Fig. 3 J).......... Heraeus caliginosus Slater & Baranowski, 1994
– Labium extending only to mesocoxae, at most reaching metacoxae; posterior pronotal lobe without four distinct spots...............................................................................................................................................8
8. Apical third of metafemur contrastingly dark brown (Fig. 11 A, B)..............................................................9
– Metafemur uniformly pale or brown, if paler basally, not strongly contrasting............................................10
9. Posterior pronotal lobe brown, mottled with paler areas, sometimes forming irregular lines; profemur dark brown with apex usually paler (Central America and Colombia; Figs 3 F, 11A) .............. Heraeus guttatus (Dallas, 1852)
– Posterior pronotal lobe uniformly dark, with two short yellow lines; profemur uniformly dark brown (Antigua, Dominican Republic, and Haiti; Figs 3 G, 11B)............................................... Heraeus hollyae Baranowski, 2005
10.Metafemur reddish brown; head smooth; distiflagellomere uniformly brown; males without small spines on protibia and mesofemur (Dominican Republic; Fig. 3 K).............................................. Heraeus dominicanus sp. nov.
– Metafemur pale; head rugose; distiflagellomere with pale band; males with small spines on protibia and mesofemur. .................................................................................................................................................... 11
11. Head, pronotum, and most of hemelytra anterior to distal white spot reddish or cinnamon brown; legs brownish yellow (USA and West Indies; Fig. 3 H) ....................................... Heraeus triguttatus (Guérin-Méneville, 1857)
– Head and anterior pronotal lobe dark brown, posterior pronotal lobe and hemelytra brown anterior to distal spot, mottled with yellow; legs uniformly pale, almost white (Dominican Republic; Fig. 3 I)...................................... .................................................................................................................... Heraeus steineri sp. nov.
12.Profemur pale brown with dark-brown spots, if predominantly dark brown, the dorsal region clearly pale (illitus group; Figs 16, 21) ........................................................................................................................... 13
– Profemur uniformly dark brown, some species with apex paler (plebejus group; Figs 26, 33, 39) .................... 22
13.Labium long, extending to metacoxae or beyond....................................................................................14
– Labium shorter, not extending beyond mesocoxae..................................................................................15
14.Strongly setose; metatibiae densely setose, with setae two times the diameter of the segment; labium long, surpassing metacoxae, usually extending to abdominal segment II (Central and South America; Fig. 15 D).............. ................................................................................................................. Heraeus illitus Distant, 1882
– Less setose; metatibial setae less abundant and shorter; labium shorter, usually extending only to posterior margin of metasternum (Argentina, Bolivia, Brazil, French Guiana, and Peru; Fig. 15 G)......... Heraeus similis sp. nov.
15.Basiflagellomere thickened distally, diameter subequal to or greater than diameter of scape.........................16
– Basiflagellomere slender, diameter less than diameter of scape................................................................ 17
16.Pronotum with a pale inverted V-shaped mark; outer margin of clavus pale (Panama; Fig. 15 I)........................ ................................................................................................................ Heraeus splendens sp. nov.
– Pronotum and hemelytra uniformly pale brown, with evenly scattered brown-stained punctures, except for distal white marks (Ecuador; Fig. 15 B)................................................................... Heraeus antennalis sp. nov.
17.Distiflagellomere uniformly brown (Argentina and Brazil; Fig. 15 C) .............. Heraeus chamamecinus sp. nov.
– Distiflagellomere with a distinct white band sub-basally.........................................................................18
18.Profemur with a distinct subapical brown band (Figs 16 A, 21B)...............................................................19
– Profemur without a subapical brown band ...........................................................................................20
19.Brown dark band on profemur narrow and contrasting (Fig. 16 A); apex of pygophore (Fig. 17 A–D) rounded, without a small dorsally directed protuberance; aedeagus with lobes sclerotized with a few spines distally (Central America, from Mexico to Panama; Fig. 15 A).................................................................. Heraeus annulatus sp. nov.
– Brown band on profemur broader and diffuse (Fig. 21 B); pygophore with a small dorsally directed protuberance near apex (Fig. 20 N); aedeagus with spines on lobes and laterally to the ejaculatory reservoir (Brazil and Ecuador; Fig. 15 H)...................................................................................................... Heraeus spinosus sp. nov.
20.Profemur irregularly pigmented with brown, metafemoral band subapical, processus gonopori long and slender (Panama; Figs 15 F, 16F) ........................................................................................ Heraeus panamaensis sp. nov.
– Profemur paler dorsally, metafemoral band apical (a few specimens of H. inca sp. nov. show a dark band on metafemur instead of being darker distally), processus gonopori long, widening towards apex (Fig. 21 D).........................21
21.Pronotum usually without contrasting dark areas laterally; dorsal anterior margin of aperture of pygophore rounded (Fig. 23 E); aedeagus with spines restricted to unsclerotized lobes (Fig. 8 I) (Ecuador; Fig. 15 J).......................... ................................................................................................................... Heraeus tiputini sp. nov.
– Pronotum usually with contrasting dark areas laterally; dorsal anterior margin of aperture of pygophore subquadrangular (Fig. 20 A); aedeagus with two pairs of lobes, the anterior lobe short, not sclerotized, and with spines; the posterior lobe large, strongly sclerotized posteriorly, with a row of spines (Peru; Fig. 15 E)............................................ ........................................................................................................................ Heraeus inca sp. nov.
22.Distiflagellomere without a pale band..................................................................................................23
– Distiflagellomere with a pale band......................................................................................................25
23.Antenna uniformly brown; dorsal margin of pygophore in lateral view almost straight and slightly declivent posteriorly (Fig. 40 F) (Galapagos Islands; Fig. 36 E).............................................. Heraeus pacificus Barber, 1925
– Antenna brown with apex of basiflagellomere and distiflagellomere darkened; dorsal margin of pygophore in lateral view sinuate and more declivent posteriorly (Fig. 41 B, F) ....................................................................... 24
24.Small species, less than 5.00 mm; subapical pale spot on corium well defined (West Indies; Fig. 36 I)................. ............................................................................................................ Heraeus pulchellus Barber, 1954
– Larger species, over 5.25 mm; subapical pale spot irregular, not well defined (USA and Central America; Fig. 36 H). Heraeus plebejus Stål, 1874
25.Labial segment IV extending only to mesocoxae .................................................................................... 26
– Labial segment IV extending to at least metacoxae................................................................................32
26.Metafemoral band apical (Figs 25 B, 26B)............................................................ Heraeus apicalis sp. nov.
– Metafemoral band subapical .............................................................................................................. 27
27.Pale band on distiflagellomere short, about one-quarter of segment length (Argentina and Bolivia; Fig. 25 F) ...... .............................................................................................................. Heraeus brevirostris sp. nov.
– Pale band on distiflagellomere longer, occupying at least one-third of segment length..................................28
28.Membranae with whitish apical spot...................................................................................................29
– Membranae without whitish apical spot...............................................................................................31
29.Anterior margin of dorsal aperture of pygophore rounded (Fig. 34 I) (Costa Rica and Mexico; Fig. 36 B) .............. ............................................................................................................... Heraeus mexicanus sp. nov.
– Anterior margin of dorsal aperture of pygophore almost straight (Figs 27 I, 40A).........................................30
30.Dark species; head blackish; posterior lobe of pronotum dark brown, with four longitudinal pale stripes (Nicaragua and Costa Rica; Fig. 36 D).................................................................. Heraeus nicaraguensis sp. nov.
– Pale species; head dark reddish brown; posterior lobe of pronotum pale brown with four difuse longitudinal pale stripes (Brazil; Fig. 25 C)................................................................................ Heraeus bahiensis sp. nov.
31.Apex of pygophore broadly rounded posteriorly in dorsal view (Fig. 32 A); outer projection of paramere not conspicuous (Fig. 32 C, D) (Bolivia; Fig. 25 E)......................................................... Heraeus bolivianus sp. nov.
– Apex of pygophore acutely rounded posteriorly in dorsal view (Fig. 34 E); outer projection of paramere conspicuous (Fig. 34 G, H) (Central America; Fig. 36 A)............................................... Heraeus mesoamericanus sp. nov.
32.Ocelli located anterior to an imaginary line passing through posterior margin of eyes; pygophore with a conspicuous protuberance at posterior margin pointed dorsally (Fig. 27 B) (Brazil; Fig. 25 A)......................................... ............................................................................................................... Heraeus alvarengai sp. nov.
– Ocelli located at level of an imaginary line passing through the posterior margin of eyes; pygophore without a dorsally directed protuberance at posterior margin, but if present, smaller and indistinct.............................33
33.Pale band on distiflagellomere narrow, less than one-third length of segment.............................................34
– Pale band on distiflagellomere broader, more than one-third length of segment .......................................... 36
34.Metafemur with a broad, subapical, dark band (Fig. 39 C); pygophore in lateral view abruptly declivent posteriorly (Fig. 40 J) (Bolivia and Brazil; Fig. 36 F)....................................................... Heraeus pallidinervis sp. nov.
– Metafemur with a narrower subapical and well-defined dark band (Figs 33C, 39D); pygophore in lateral view slightly declivent posteriorly (Figs 34 B, 40N)................................................................................................35
35.Inner projections of pygophore subquadrangular and slightly projecting posteriorly (Fig. 40 M) (Argentina, Bolivia, and Paraguay; Fig. 36 G)...................................................................................... Heraeus penai sp. nov.
– Inner projections of pygophore narrow, elongate, and strongly projecting posteriorly (Fig. 34 A) (Ecuador; Fig. 25 I).............................................................................................................. Heraeus loja sp. nov.
36.Metafemur with a narrow subapical dark band (Fig. 26 D); pygophore projecting posteriorly and dorsally directed (Fig. 27 N) (Belize and Mexico; Fig. 25 D)........................................................ Heraeus baranowski sp. nov.
– Metafemur with a broader subapical dark band (Fig. 33 A, B, F); pygophore not projecting posteriorly nor pointed dorsally (Figs 32 J, N, 34N)................................................................................................................37
37.Small species, less than 5 mm long; pygophore with inner projections of dorsal aperture subquadrangular (Fig. 32 I) (Dominican Republic and Haiti; Fig. 25 G) ................................... Heraeus concolor Slater & Baranowski, 1994
– Larger species, over 5 mm long; pygophore with inner projections of dorsal aperture elongate, not subquadrangular (Figs 32 M, 34M) .............................................................................................................................. 38
38.Pygophore rounded; with inner projections of dorsal aperture subtriangular, not well delimited anteriorly (Fig. 32 M); parameres with inner projection narrow (Fig. 32 O, P) (Ecuador; Fig. 25 H) ........ Heraeus ecuatorianus sp. nov.
– Pygophore declivent and truncate posteriorly in lateral view (Fig. 34 N); inner projections of dorsal aperture elongate, well delimited anteriorly (Fig. 34 M); parameres with inner projection broad (Fig. 34 O, P) (Panama; Fig. 36 C).................................................................................................... Heraeus morganae sp. nov.
description: Published as part of Pablo M. Dellapé, María C. Melo & Thomas J. Henry, 2016, A phylogenetic revision of the true bug genus Heraeus (Hemiptera: Rhyparochromidae: Myodochini), with the description of two new genera and 30 new species, pp. 29-134 in Zoological Journal of the Linnean Society 177 (177) on pages 38-40, DOI: 10.1111/zoj.12362, http://zenodo.org/record/269748