Heraeus

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, Zoological Journal of the Linnean Society 177 (177), pp. 29-134 : 38-40

publication ID	https://doi.org/ 10.1111/zoj.12362
publication LSID	lsid:zoobank.org:pub:8A961A76-E33C-4171-B401-0B516B62FA6D
DOI	https://doi.org/10.5281/zenodo.5611107
persistent identifier	https://treatment.plazi.org/id/6F64DB1B-5E49-FFA4-FDE4-FF05FA4AFC95
treatment provided by	Plazi
scientific name	Heraeus
status

Treatment

KEY TO THE SPECIES OF HERAEUS View in CoL

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 View Figure 3 A–E); head smooth ( coquilletti View in CoL group) ................................................... 2

– Hemelytra more heterogeneously coloured, often with a large pale spot at apex of corium ( Fig. 3 View Figure 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 View Figure 3 D) ... ..................................................................................................................... Heraeus itzelae View in CoL sp. nov.

– Distiflagellomere yellowish brown, without a pale band; hemelytra with long, erect setae (Honduras and Mexico; Fig. 3 View Figure 3 E).......................................................................................................... Heraeus setosus View in CoL sp. nov.

4. Head, pronotum, and hemelytra shiny (Mexico and USA; Fig. 3 View Figure 3 B).................... Heraeus coquilletti Barber, 1914 View in CoL

– 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 View Figure 3 C) ................................................... Heraeus costalis View in CoL sp. nov.

– General colouration cinnamon brown; distiflagellomere uniformly brown; profemur pale yellowish brown (Mexico and USA; Fig. 3 View Figure 3 A)............................................................................... Heraeus cinnamomeus Barber, 1948 View in CoL

6. Hemelytron with a distinct, white, subapical spot on corium and a white mark on apex of membrane; species usually glabrous ( guttatus View in CoL and caliginosus View in CoL groups; Fig. 3 View Figure 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 View Figure 15 , 25 View Figure 25 , 36 View Figure 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 View Figure 3 J).......... Heraeus caliginosus Slater & Baranowski, 1994 View in CoL

– 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 View Figure 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 View Figure 3 F, 11A) .............. Heraeus guttatus ( Dallas, 1852) View in CoL

– Posterior pronotal lobe uniformly dark, with two short yellow lines; profemur uniformly dark brown (Antigua, Dominican Republic, and Haiti; Figs 3 View Figure 3 G, 11B)............................................... Heraeus hollyae Baranowski, 2005 View in CoL

10.Metafemur reddish brown; head smooth; distiflagellomere uniformly brown; males without small spines on protibia and mesofemur (Dominican Republic; Fig. 3 View Figure 3 K).............................................. Heraeus dominicanus View in CoL 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 View Figure 3 H) ....................................... Heraeus triguttatus ( Guérin-Méneville, 1857) View in CoL

– 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 View Figure 3 I)...................................... .................................................................................................................... Heraeus steineri View in CoL sp. nov.

12.Profemur pale brown with dark-brown spots, if predominantly dark brown, the dorsal region clearly pale ( illitus View in CoL group; Figs 16 View Figure 16 , 21 View Figure 21 ) ........................................................................................................................... 13

– Profemur uniformly dark brown, some species with apex paler ( plebejus View in CoL group; Figs 26 View Figure 26 , 33 View Figure 33 , 39 View Figure 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 View Figure 15 D).............. ................................................................................................................. Heraeus illitus Distant, 1882 View in CoL

– 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 View Figure 15 G)......... Heraeus similis View in CoL 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 View Figure 15 I)........................ ................................................................................................................ Heraeus splendens View in CoL sp. nov.

– Pronotum and hemelytra uniformly pale brown, with evenly scattered brown-stained punctures, except for distal white marks (Ecuador; Fig. 15 View Figure 15 B)................................................................... Heraeus antennalis View in CoL sp. nov.

17.Distiflagellomere uniformly brown (Argentina and Brazil; Fig. 15 View Figure 15 C) .............. Heraeus chamamecinus View in CoL sp. nov.

– Distiflagellomere with a distinct white band sub-basally.........................................................................18

18.Profemur with a distinct subapical brown band ( Figs 16 View Figure 16 A, 21B)...............................................................19

– Profemur without a subapical brown band ...........................................................................................20

19.Brown dark band on profemur narrow and contrasting ( Fig. 16 View Figure 16 A); apex of pygophore ( Fig. 17 View Figure 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 View Figure 15 A).................................................................. Heraeus annulatus View in CoL sp. nov.

– Brown band on profemur broader and diffuse ( Fig. 21 View Figure 21 B); pygophore with a small dorsally directed protuberance near apex ( Fig. 20 View Figure 20 N); aedeagus with spines on lobes and laterally to the ejaculatory reservoir (Brazil and Ecuador; Fig. 15 View Figure 15 H)...................................................................................................... Heraeus spinosus View in CoL sp. nov.

20.Profemur irregularly pigmented with brown, metafemoral band subapical, processus gonopori long and slender (Panama; Figs 15 View Figure 15 F, 16F) ........................................................................................ Heraeus panamaensis View in CoL sp. nov.

– Profemur paler dorsally, metafemoral band apical (a few specimens of H. inca View in CoL sp. nov. show a dark band on metafemur instead of being darker distally), processus gonopori long, widening towards apex ( Fig. 21 View Figure 21 D).........................21

21.Pronotum usually without contrasting dark areas laterally; dorsal anterior margin of aperture of pygophore rounded ( Fig. 23 View Figure 23 E); aedeagus with spines restricted to unsclerotized lobes ( Fig. 8 View Figure 8 I) (Ecuador; Fig. 15 View Figure 15 J).......................... ................................................................................................................... Heraeus tiputini View in CoL sp. nov.

– Pronotum usually with contrasting dark areas laterally; dorsal anterior margin of aperture of pygophore subquadrangular ( Fig. 20 View Figure 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 View Figure 15 E)............................................ ........................................................................................................................ Heraeus inca View in CoL 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 View Figure 40 F) (Galapagos Islands; Fig. 36 View Figure 36 E).............................................. Heraeus pacificus Barber, 1925 View in CoL

– Antenna brown with apex of basiflagellomere and distiflagellomere darkened; dorsal margin of pygophore in lateral view sinuate and more declivent posteriorly ( Fig. 41 View Figure 41 B, F) ....................................................................... 24

24.Small species, less than 5.00 mm; subapical pale spot on corium well defined (West Indies; Fig. 36 View Figure 36 I)................. ............................................................................................................ Heraeus pulchellus Barber, 1954 View in CoL

– Larger species, over 5.25 mm; subapical pale spot irregular, not well defined (USA and Central America; Fig. 36 View Figure 36 H). Heraeus plebejus Stål, 1874 View in CoL

25.Labial segment IV extending only to mesocoxae .................................................................................... 26

– Labial segment IV extending to at least metacoxae................................................................................32

26.Metafemoral band apical ( Figs 25 View Figure 25 B, 26B)............................................................ Heraeus apicalis View in CoL sp. nov.

– Metafemoral band subapical .............................................................................................................. 27

27.Pale band on distiflagellomere short, about one-quarter of segment length (Argentina and Bolivia; Fig. 25 View Figure 25 F) ...... .............................................................................................................. Heraeus brevirostris View in CoL 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 View Figure 34 I) (Costa Rica and Mexico; Fig. 36 View Figure 36 B) .............. ............................................................................................................... Heraeus mexicanus View in CoL sp. nov.

– Anterior margin of dorsal aperture of pygophore almost straight ( Figs 27 View Figure 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 View Figure 36 D).................................................................. Heraeus nicaraguensis View in CoL sp. nov.

– Pale species; head dark reddish brown; posterior lobe of pronotum pale brown with four difuse longitudinal pale stripes (Brazil; Fig. 25 View Figure 25 C)................................................................................ Heraeus bahiensis View in CoL sp. nov.

31.Apex of pygophore broadly rounded posteriorly in dorsal view ( Fig. 32 View Figure 32 A); outer projection of paramere not conspicuous ( Fig. 32 View Figure 32 C, D) (Bolivia; Fig. 25 View Figure 25 E)......................................................... Heraeus bolivianus View in CoL sp. nov.

– Apex of pygophore acutely rounded posteriorly in dorsal view ( Fig. 34 View Figure 34 E); outer projection of paramere conspicuous ( Fig. 34 View Figure 34 G, H) (Central America; Fig. 36 View Figure 36 A)............................................... Heraeus mesoamericanus View in CoL 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 View Figure 27 B) (Brazil; Fig. 25 View Figure 25 A)......................................... ............................................................................................................... Heraeus alvarengai View in CoL 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 View Figure 39 C); pygophore in lateral view abruptly declivent posteriorly ( Fig. 40 View Figure 40 J) (Bolivia and Brazil; Fig. 36 View Figure 36 F)....................................................... Heraeus pallidinervis View in CoL sp. nov.

– Metafemur with a narrower subapical and well-defined dark band (Figs 33C, 39D); pygophore in lateral view slightly declivent posteriorly ( Figs 34 View Figure 34 B, 40N)................................................................................................35

35.Inner projections of pygophore subquadrangular and slightly projecting posteriorly ( Fig. 40 View Figure 40 M) (Argentina, Bolivia, and Paraguay; Fig. 36 View Figure 36 G)...................................................................................... Heraeus penai View in CoL sp. nov.

– Inner projections of pygophore narrow, elongate, and strongly projecting posteriorly ( Fig. 34 View Figure 34 A) (Ecuador; Fig. 25 View Figure 25 I).............................................................................................................. Heraeus loja View in CoL sp. nov.

36.Metafemur with a narrow subapical dark band ( Fig. 26 View Figure 26 D); pygophore projecting posteriorly and dorsally directed ( Fig. 27 View Figure 27 N) (Belize and Mexico; Fig. 25 View Figure 25 D)........................................................ Heraeus baranowski View in CoL sp. nov.

– Metafemur with a broader subapical dark band ( Fig. 33 View Figure 33 A, B, F); pygophore not projecting posteriorly nor pointed dorsally ( Figs 32 View Figure 32 J, N, 34N)................................................................................................................37

37.Small species, less than 5 mm long; pygophore with inner projections of dorsal aperture subquadrangular ( Fig. 32 View Figure 32 I) (Dominican Republic and Haiti; Fig. 25 View Figure 25 G) ................................... Heraeus concolor Slater & Baranowski, 1994 View in CoL

– Larger species, over 5 mm long; pygophore with inner projections of dorsal aperture elongate, not subquadrangular ( Figs 32 View Figure 32 M, 34M) .............................................................................................................................. 38

38.Pygophore rounded; with inner projections of dorsal aperture subtriangular, not well delimited anteriorly ( Fig. 32 View Figure 32 M); parameres with inner projection narrow ( Fig. 32 View Figure 32 O, P) (Ecuador; Fig. 25 View Figure 25 H) ........ Heraeus ecuatorianus View in CoL sp. nov.

– Pygophore declivent and truncate posteriorly in lateral view ( Fig. 34 View Figure 34 N); inner projections of dorsal aperture elongate, well delimited anteriorly ( Fig. 34 View Figure 34 M); parameres with inner projection broad ( Fig. 34 View Figure 34 O, P) (Panama; Fig. 36 View Figure 36 C).................................................................................................... Heraeus morganae View in CoL sp. nov.