A phylogenetic revision of the true bug genus Heraeus (Hemiptera: Rhyparochromidae: Myodochini), with the description of two new genera and 30 new species Author Dellapé, Pablo M. Author Melo, María C. Author Henry, Thomas J. text Zoological Journal of the Linnean Society 2016 2016-04-17 177 1 29 134 http://dx.doi.org/10.1111/zoj.12362 journal article 10.1111/zoj.12362 0024-4082 10109299 8A961A76-E33C-4171-B401-0B516B62FA6D HERAEUS COQUILLETTI BARBER, 1914 ( FIGS 3B , 4D–E , 5B , 6E–H , 7 , 8A ) Heraeus coquilletti Barber, 1914a: 165–166 (in part); Van Duzee, 1916: 21 (in part); Van Duzee, 1917: 179 (in part); Blatchley, 1934: 8 (in part); Torre Bueno, 1946: 70 (in part); Slater, 1964: 1082 ; Harrington, 1980: 108 ; Ashlock & Slater, 1988: 227–228 ; Slater & Brailovsky, 2000: 332 . Heraeus nitens Van Duzee, 1914: 8–9 (synonymized by Van Duzee, 1916 ). Diagnosis Heraeus coquilletti can be easily distinguished from all other species of the genus by the shiny head, pronotum, and hemelytra with minute setae. In addition, the distiflagellomeres and hemelytra are uniformly brown and the aedeagus has minute spines on conjunctiva and vesica. Redescription Body shiny, with minute setae on head, pronotum, scutellum and hemelytra ( Fig. 3B ). Head: Strongly convex dorsally, orange–brown, smooth, with sparse short recumbent setae. Head transversely rugose ventrally. Eyes slightly protruding, small (larger in specimens from Mexico ), not surpassing the dorsal margin of head in lateral view. Ocelli posteri- or to an imaginary line passing through posterior border of eyes. Labium pale brown with short erect setae, extending to mesocoxae. Antennal tubercles divergent. Antennae pale brown, with short recumbent and scattered erect setae, basiflagellomere and distiflagellomere darker, with more abundant setae. Figure 3. Habitus dorsal: A, Heraeus cinnamomeus Barber, 1948 ; B, Heraeus coquilletti Barber, 1914 ; C, Heraeus costalis sp. nov. ; D, Heraeus itzelae sp. nov. ; E, Heraeus setosus sp. nov. ; F, Heraeus guttatus ( Dallas, 1852 ) . G, Heraeus hollyae Baranowski, 2005 . H, Heraeus triguttatus ( Guérin-Méneville, 1857 ) ; I, Heraeus steineri sp. nov. ; J, Heraeus caliginosus Slater & Baranowski, 1994 ; K, Heraeus dominicanus sp. nov. Thorax: Pronotum dark orange–brown, smooth ( Fig. 4D ); posterior lobe paler than anterior lobe; punctate, punctures more conspicuous and abundant on posterior lobe; anterior lobe with sparse, minute, erect setae. Pleurae orange–brown. Evaporative area short. Scutellum orange–brown, punctate. Hemelytra shiny, smooth ( Fig. 4E ), pale brown, with abundant punctures. Membrane smoky pale brown. Legs: Coxae orange–brown, femora, tibiae, and tarsi pale brown, except apex of each tibia darker, meso- and metafemur slightly darkened apically ( Fig. 5B ). Male profemur strongly enlarged with two rows of spines along ventral region, four spines from the apex on posterior row longer and curved anteriorly, with short and sparse setae. Protibia with numerous small tubercles, metatibia with spiniform setae. Figure 4. Heraeus cinnamomeus Barber, 1948 : A, head and pronotal collar, lateral view; B, detail of pruinosity on anterior pronotal lobe; C, detail of pruinosity of part of scutellum and clavus. Heraeus coquilletti Barber, 1914 : D, detail of texture of anterior pronotal; E, detail of texture of part of scutellum and clavus. Abbreviations: cl, clavus, co, corium; sc, scutellum. Abdomen: Orange–brown with short sparse recumbent setae. Male genitalia: Pygophore ( Fig. 6E, F ) rounded, anterior margin of dorsal aperture rounded, inner projections subrectangular. Parameres: Figure 6 (G, H). Aedeagus ( Fig. 8A ): conjunctiva with a few minute spines laterally; lobes of vesica slightly sclerotized, with a few minute spines laterally and distally; processus gonopori long and slender. Distribution Mexico and USA ( Fig. 7 ). Remarks Figure 5. Legs: A, Heraeus cinnamomeus Barber, 1948 ; B, Heraeus coquilletti Barber, 1914 ; C, Heraeus costalis sp. nov. ; D, Heraeus itzelae sp. nov. ; E, Heraeus setosus sp. nov. ; F, Heraeus caliginosus Slater & Baranowski, 1994 ; G, Heraeus dominicanus sp. nov. Barber (1914a) based his original description on two males and two females from Standford University, California (Nathan Banks Collection), and a female from Brownsville, Texas , USA , that was referred to as H. cinnamomeus by Barber (1948) . Type material examined To ensure nomenclatural stability, we select a male syntype in the USNM collection with the following labels as the lectotype of H. coquilletti : Label 1, ‘Stan[ford] U[.], Cal[.], 24 Dec 1909 ’; 2 (red) ‘Cotype’; 3, ‘H.G. Barber Collection’; 4 (handwritten), ‘ Heraeus coquilletti Barber [,] ♂ [,] Cotype’; 5 (red, here added), Lectotype : ♂ , Heraeus coquilletti Barber , desig. by Dellapé, Melo, & Henry’. A female labelled as co-type, also in the USNM collection, and a male (missing the head) in the AMNH collection with the same locality data are considered paralectotypes . Additional material studied MEXICO : Baja California : 1♀ , Santo Tomás , 8-VII- 1953 , W.J. & J.W. Gertsch ( AMNH ) ; 1♀ , Mexico , Brassica sp. , 18-II-2003 , San Diego 030359, Miami Port , March 2007 ( USNM ) ; Sinaloa : 1♂ , 40 mi. N Mazatlan , 27-VII-1952 , at light, J.D. Lattin ( AMNH ) ; 8♂ , 10♀ , 26 mi. N Pericos , 13-VIII-1960 , P.H. Arnaud Jr , E.S. Ross & D.C. Reutz ( CAS ) ; 1♂ , 1♀ , Mazatlan , 27-VI-1918 , Venedio, J.A . Kusche , pres. by B. Preston Clark ( CAS ) ; 1♂ , 2♀ , 1-VI-1918 , J.A. Kusche, pres. by B. Preston Figure 6. Male genitalia. Heraeus cinnamomeus Barber, 1948 : A, pygophore, dorsal view; B, pygophore, lateral view; C, right paramere, inner view; D, right paramere, external view. Heraeus coquilletti Barber, 1914 : E, pygophore, dorsal view; F, pygophore, lateral view; G, right paramere, inner view; H, right paramere, external view. Heraeus costalis sp. nov. : I, pygophore, dorsal view; J, pygophore, lateral view; K, right paramere, inner view; L, right paramere, external view. Heraeus itzelae sp. nov. : M, pygophore, dorsal view; N, pygophore, lateral view; O, right paramere, inner view; P, right paramere, external view. Figure 7. Distributional map of Heraeus cinnamomeus Barber, 1948 , Heraeus coquilletti Barber, 1914 , Heraeus costalis sp. nov. , Heraeus itzelae sp. nov. , and Heraeus setosus sp. nov. Clark ( CAS ); 1♂ , 5-VI-1918 , J.A. Kusche, pres. by B. Preston Clark ( CAS ); 2♂ , Presidio Riv., 26-IX- [19]18, J.A. Kusche ( USNM ). USA : Arizona : one without abdomen, Phoenix ( USNM ) ; California : 2♂ , Claremont , Baker ( CAS ) ; 1♀ , San Joaquin Co. , 9 mi. E Stockton , 10-VIII-1971 , Chenopodium , D. Shepard , ( AMNH ) ; 1♂ , 2♀ , Oakland , Alameda Co. , 25-II-1906 , van Dyke ( CAS ) ; 1♀ , Alameda Co. , van Dyke ( CAS ) ; 1♀ , van Dyke ( CAS ) ; 1♂ , Burbank , 22-II-1930 , C.H. Hicks , ( USNM ) ; 1♀ , 24-VIII-1930 ( USNM ) ; 1♀ , Leona Hgts , Alameda , Aug., J.C . Bradley ( CAS ) ; 1♂ , Alameda , Nov. , Koebele ( CAS ) ; 1♂ , 1♀ , S Sonoma Co. , 18-II-1911 , van Dyke ( CAS ) ; 1♀ , Sonoma Co. , VI-1919 , J.R. de la Torre Bueno ( CAS ) ; 1♂ , 1♀ , Sonoma Co. , C. Olsen ( AMNH ) ; 2♂ , Berkeley , 15-I-1922 , pres. by E.C. van Dyke ( CAS ) ; 1♀ , Redwood City , 1-I-1943 , P.H. Arnaud ( CAS ) ; 1♂ , 1♀ , San Mateo Co. , 8-I-1944 , P.H. Arnaud , 20– 161 ( CAS ) ; 1♂ , 8-I-1944 , P.H. Arnaud , 20–161 ( USNM ) ; 1♀ , Oroville , 13-VII-1926 , H.H. Kelfer ( CAS ) ; 1♀ , Fairfax , 9-III-1919 , E.P. Van Duzee ( CAS ) ; 1♂ , Redding , Shasta Co. , 31-VII-1947 , elev. 500, H.P. Chandler ( CAS ) ; 6♂ , 3♀ , San Luis Obispo , 28- IV-1919 , E.P. van Duzee ( CAS ) ; 1♂ , Morro beach, 6-IV- 1928 , pres. by E.R. Lesch ( CAS ) ; 1♂ , 2♀ , Morro , S.L. Ob. Co. , 20-XII-1928 , pres. by E.C. van Dyke ( CAS ) ; 2♀ , Romero Canyon , San Ysidro , 3-I-[19]11, W.M. Wheeler & H.M. Parshley ( CAS ) ; 1♀ , 3-I- [19]11, W.M. Wheeler & H.M. Parshley , V.D. no. 529 ( CAS ) ; 1♂ , Ojai , Ventura Co. , 16-VI-1957 , W.E. Simonds , ( AMNH ) ; 1♀ Madera , VI-22-[19]59, C.A. Toschi ( AMNH ) ; 2♂ , 1♀ Sonoma Co. , C.E. Olsen ( AMNH ) ; 1♂ , H.G. Barber ( USNM ) ; 1♀ , Vacaville , 6-IX-[19]47, A.T. McClay ( USNM ) ; 1♀ Monterrey Co. , 4.5 mi. N Parkfield , el. abt 2000 ft , IV-2-1977 , K.W. Brown , R.E. Somerby ( AMNH ) ; 4♂ , 5♀ , Los Angeles , Coquillett ( USNM ) ; 3♂ , Los Angeles, P.R . Uhler ( USNM ) ; 3♂ , 5♀ , San Diego, 30-I, Hubbard Figure 8. Aedeagus. A, Heraeus coquilletti Barber, 1914 ; B, Heraeus costalis sp. nov. ; C, Heraeus setosus sp. nov. ; D, Heraeus caliginosus Slater & Baranowski, 1994 ; E, Heraeus guttatus ( Dallas, 1852 ) ; F, Heraeus chamamecinus sp. nov. , anterior view; G, Heraeus chamamecinus sp. nov. , lateral view; H, Heraeus illitus Distant, 1882 ; I, Heraeus tiputini sp. nov. ( USNM ); 1♀ , 11-III-[19]14, E.P. Van Duzee ( USNM ) ; New Mexico : 1♂ , Sandoval Co. , San Ysidero , IX- 1967 , Wilton Ivie ( AMNH ) .