Revision of the African shieldbug genus Afrius Stål, 1870 (Hemiptera: Heteroptera: Pentatomidae: Asopinae) Author Roell, Talita Author Lemaître, Valérie A. Author Webb, Michael D. text European Journal of Taxonomy 2019 2019-04-24 520 1 44 journal article 26982 10.5852/ejt.2019.520 de14dab5-2961-423b-867f-5a4a48d8bb3e 2650382 085164DC-B31D-4293-A037-9C31940E24CD Afrius ( Afrius ) purpureus ( Westwood, 1837 ) Figs 1 , 8–11 Pentatoma yolofa Guérin-Méneville, 1831: 55 , fig. 2 and legend (syn. Stål 1870: 44 ). Pentatoma purpurea Westwood, 1837: 43 . Asopus figuratus Germar, 1838: 185 –186 (syn. Schouteden 1905: 147 , as a var. of Canthecona purpurea ). Canthecona caerulea Dallas, 1851: 89 (syn. Stål 1862: 496 , as var. of Canthecona Yolofa Guérin ; syn. Stål 1870: 44 ). Canthecona marginella Dallas, 1851: 89 (syn. Stål 1862: 496 , as var. of Canthecona Yolofa Guérin ; syn. Stål 1870: 44 ). Canthecona marmorata Dallas, 1851: 90 (synonymy with Canthecona purpurea suspected by Schouteden 1905a: 150). syn. nov. Canthecona annulipes Dallas, 1851: 90 –91 (syn. Schouteden 1905a: 149–150, with Canthecona marmorata ; synonymy with Canthecona purpurea suspected by Schouteden 1905a: 150). syn. nov. Canthecona miniatescens Stål, 1854: 213 (syn. Stål 1864: 66 –67 , with Canthecona figurata , as var. b.; syn. Kirkaldy 1909: 10 , with Afrius purpureus , as a var.). Afrius rubromarginatus Bergroth, 1903: 289 . syn. nov. Pentatoma ( Eurydema ) yolofa – Laporte 1833: 61 . Canthecona Yolofa – Amyot & Serville 1843: 82 . — Dallas 1851: 89 . — Stål 1864: 67 –68 . — Larousse 1890: 724 [without any capital letters]. Cimex ( Pentatoma ) yolofa – Guérin-Méneville 1844: 344 [description] Asopus figuratus – Herrich-Schaffer 1844: 113 , fig. 710. Canthecona figurata – Stål 1864: 66 –67 . — Gerstaecker 1892: 345 . Cimex ( Afrius ) figuratus – Stål 1870: 44 . Cimex (Afrius) purpureus – Stål 1870: 44 . — Distant 1884: 459 . Cimex marmoratus – Stål 1870: 46 . Cimex annulipes – Stål 1870: 46 . Canthecona Ylofa (incorrect subsequent spelling) – Wallengren 1875: 133 . Afrius purpureus – Reuter 1882: 9 . — Lethierry & Severin 1893: 214 . — Schouteden 1907a: 51–52 ; 1909: 64 ; 1910: 91 ; 1913a: 190 ; 1913b: 324 ; 1963: 399 ; 1964: 95 ; 1972: 106 . — Kirkaldy 1909: 10 . — Vuillet & Vuillet 1911: 277 ; 1912: 445 . — Hollrung 1912: 280 . — Jeannel 1913: 97 . — Zacher 1921: 139 . — Lehmann 1922: 129 . — Hesse 1925: 39 . — Carpenter 1926 : liii. — Golding 1931: 222 . — Van Heerden 1931: 131 , fig. cvii. — Hargreaves 1937: 518 . — Villiers 1949: 88 ; 1952a: 1211 ; 1952b: 82 ; 1954: 230 ; 1956: 213 . — Risbec 1950: 126 , 397, 447, 457. — Cachan 1952: 305 ; 1965: 5–32 . — Leston 1952: 895 . — Mancini 1953: 202 . — Lindberg 1958: 31 . — Le Pelley 1959: 166 . — Caswell 1962: 26 . — Girard 1969: 7 , 52. — Herting 1971: 80 ; 1976: 71 . — Gillon 1972: 351 –352 ; 1974: 219 , 241, 246, 251, 266, 270, 285, 287. — Linnavuori 1975: 124 ; 1976: 129 ; 1982: 164 . — Medler 1980: 123 . — Bourdouxhe & Jolivet 1981: 46 –48 . — Nuamah 1982: 11 . — Nonveiller 1984: 54 , 142. — Balsbaugh 1988: 276 . — Couilloud 1989: 207 –208 . — Sylvie et al. 1989: 277. — Quicke et al. 1992: 1024 . — Van Harten 1993: 247 . — Bijlmakers & Verhoek 1995: 147 , 317. — Schaefer 1996: 41 . — Jolivet 1997: 153 . — Boulard 1998: 41 . — Dolling et al. 1999: 73 . — Beenen & Hawkeswood 2004: 473 . — Tchibozo & Braet 2004: 161 . — Arechavaleta et al. 2005: 76 . — Renou 2007: 30 . — Poutouli et al. 2011: 9 , 54–56, 58, 76. — Agboton et al. 2014: 9 . — Crétenet & Gourlot 2015: 138 . Afrius purpureus var. marginella – Reuter 1882: 9 . — Kirkaldy 1909: 10 . Cimex ( Afrius ) purpureus var. marginella – Distant 1884: 459 . Cimex purpureus – Distant 1890 : LIII [with Hope as author, not Westwood]. Cimex figuratus – Distant 1892: 248 [description and illustration of unnamed variety]. Afrius ? annulipes – Lethierry & Severin 1893: 214 . http://www.palaeontos.be/13/det13.html Afrius figuratus – Lethierry & Severin 1893: 214 . — Distant 1898: 308 ; 1901: 27 . — Howard 1906: 731 . — Leston 1954: 680 (and in title). — Maitai 1958: 291 . — Le Pelley 1959: 257 ; 1968: 195 , 501. — McDonald 1966: 44 . — Cobben 1968: 116 . — Herting 1973: 84 , 85, 87. — Medler 1980: 123 . — Smith & Barfield 1982: 263 . — Scholtz & Holm 1985 : 147. — Merrett 1986: 549 . — Matanmi & Hassan 1987: 376 . — Fry 1989: 108 . — Mugo 1995: 2099 . — Schaefer 1996: 44 . — Sileshi et al. 2000: 41 , 48. Afrius ? marmoratus – Lethierry & Severin 1893: 214 . Canthecona purpurea – Schouteden 1905a: 146 ; 1905b: 15. Canthecona purpurea var. figuratus – Schouteden 1905a: 147 . Canthecona marmorata – Schouteden 1905a: 149 –150. Canthecona rubromarginata – Schouteden 1905a: 150 –151. Afrius marmoratus – Schouteden 1907a: 51 . — Kirkaldy 1909: 10 . — Thomas 1994: 151 . — Robertson 2009: 22 . Afrius rubromarginatus – Schouteden 1907a: 52 . — Bergroth 1908: 182 . — Kirkaldy 1909: 10 . — Thomas 1994: 151 . — Maldès & Pluot-Sigwalt 2004: 20 . — Robertson 2009: 22 . — Krüger & Deckert 2016: 46 . Afrius purpureus var. figuratus – Schouteden 1907a: 51–52 ; 1909: 64 . — Mancini 1937: 43 . — Le Pelley 1959: 54 . Afrius purpureus var. caerulea – Kirkaldy 1909: 10 . Afrius purpureus var. figurata – Kirkaldy 1909: 10 . Afrius purpureus var. miniatescens – Kirkaldy 1909: 10 . Afrius yolofus – Dupuis 1952: 454 . — Leston 1954: 680 (in a note). — Thomas 1994: 151 . — Maldès & Pluot-Sigwalt 2004: 20 . — Rebagliati et al. 2005: 201 . — Kerisew 2011: 91 . Afrius purpureus purpureus – Linnavuori 1989: 13 . Afrius purpureus figuratus – Linnavuori 1989: 12 –13 [with unnamed variety]. Afrius yolofa – Sileshi et al. 2001: 289 ; 2004: 6 , 18. — Kerzhner et al. 2004: 18 . — Rider 2006: 234 . — Robertson 2009: 21 –22 . — Matesco et al. 2014: 352 . Types examined AFRICA • Pentatoma purpurea Westwood, 1837 ; syntype ♀ ; labels: “Type Hem: 242 PENTATOMA PURPUREA WESTWOOD HOPE DEPT . OXFORD”, “ TYPE = WEST . ( HOPE ) C. Hemipt. 1837 Part. I. page 43 Distant, P.Z.S., 1900, p. 807–825.”, “Africa”, “ Afrius purpureus Westw. ”; Fig. 8 E–F ; OUMNH . “EAST AFRICA” • Afrius rubromarginatus Bergroth, 1903 ; syntype ♀ ; labels: “Type Hem: 704 AFRIUS RUBROMARGINATUS BERGROTH . HOPE DEPT . OXFORD”, “ Afrius rubro = marginatus Bergr.”, “Probably TANGANYIKA”, “Ann. Soc. Ent. Belg. 47: 289”, “Afriq Or”, “3/”, “ TYPE ”; Fig. 8 O–P ; OUMNH . SIERRA LEONE • Canthecona marginella Dallas, 1851 ; syntype ³; labels: blue-margined syntype disc label, red-margined type disc label, “87a”, “ Canthecona marginella identified by Dallas”, “a”, “ NHMUK 010592170 ”; Fig. 8 I–J ; NHMUK 0 10592170 . SOUTH AFRICA • Asopus figuratus Germar, 1838 ; syntype ♀ ; labels: “ figuratus Germ. Promont. b. sp. Collect. Germ.”, “ Afrius figuratus (Germ) ”, “7968”, “Typus”; Fig. 8 G–H ; MFNB . SOUTH AFRICA • Canthecona caerulea Dallas, 1851 ; syntype ³; labels: blue-margined syntype disc label, red-margined type disc label, “ 40 6 26 329”, “ Canthecona caerulea identified by Dallas”, “a”, “ NHMUK 010592171 ”; Fig. 8 A–C ; NHMUK 0 10592171 . SOUTH AFRICA • Canthecona marmorata Dallas, 1851 ; syntype ³; labels: blue-margined syntype disc label, red-margined type disc label, “ Int. S. Africa / 4319”, “3. Canthecona marmorata ,”, “a”, “ NHMUK 010592164 ”; Fig. 8 K–L ; NHMUK 0 10592164 . SOUTH AFRICA • Canthecona annulipes Dallas, 1851 ; syntype ♀ ; labels: blue-margined syntype disc label, “Int. Africa / 4319”, “ Canthecona figurata Walker’s catal.”, “a”, “ NHMUK 010747805 ”; Fig. 8 M–N ; NHMUK 0 10747805 . SOUTH AFRICA • Canthecona miniatescens Stål, 1854 ; syntype ♀ ; labels: “Caffraria”, “ I. Vahlb ”, “ miniatescens Stål type.”, “Typus”, “NHRS-GULI 000027293”; Fig. 8D ; examined by photo; NHRS- GULI 0 0 0 0 27293 . Diagnosis Scutellum longer than wide, postfrenal lobe enlarged; humeral pronotal angles slightly emarginated; male abdomen with setose patches on segments V and VI, parameres triangular, without two distinct lobes. Redescription BODY. Elongate oval, with variable colour patterns from yellowish and brownish with yellow or red stripes ( Fig. 8D, H, L, N ) to purple, green and blue metallic uniform colours ( Fig. 8 B–C, F, J, P). HEAD. Subrectangular, wider than long, punctured; mandibular plates equal or slightly longer than clypeus, with margin varying from straight to sinuous; antenna bearing thin setae, denser on third, fourth and fifth antennomeres, proportion of lengths of antennomeres: IV> V> II> III> I; labium robust, reaching metasternum; proportion of lengths of labiomeres: II> I> IV> III. Fig. 8. Afrius ( Afrius ) purpureus ( Westwood, 1837 ) , type specimens. A–C . Canthecona caerulea Dallas, 1851 , syntype ³, labels, dorsal and ventral habitus, respectively (NHMUK). D . Canthecona miniatescens Stål, 1854 , syntype ♀, labels and dorsal view (image courtesy of Gunvi Lindberg, NHRS). E–F . Pentatoma purpurea Westwood, 1837 , syntype ♀, labels and dorsal habitus (OUMNH). G–H . Asopus figuratus Germar, 1838 , syntype ♀, labels and dorsal habitus (MFNB). I–J . Canthecona marginella Dallas, 1851 , syntype ³, labels and dorsal habitus (NHMUK). K–L . Canthecona marmorata Dallas, 1851 , syntype ³, labels and dorsal habitus (NHMUK). M–N . Canthecona annulipes Dallas, 1851 , syntype ♀, labels and dorsal habitus (NHMUK). O–P . Afrius rubromarginatus Bergroth, 1903 , syntype ♀, labels and dorsal habitus (OUMNH). Abbreviations: ac = corium adjacent to the scutellum constriction; paVII = posterior angles of seventh abdominal segment; sc = scutellum constriction. Scale bars: 4 mm. THORAX. Pronotum hexagonal, densely punctured, twice or more as wide as long, cicatrices flat; anterior margin concave; lateral margins strongly sinuous, crenulated on anterior half ( Fig. 9 ), humeral angle slightly emarginated, anterior humeral portion varying from convex ( Fig. 9A, C ) to acute or spinose ( Fig. 9B, D ). Scutellum longer than wide, densely punctured. Corium longer than scutellum, usually not surpassing connexival segment V, densely punctured. MALE ABDOMEN. Setose patches present on male abdominal segments V and VI ( Fig. 8C ); parameres uniramous, triangular. Male MEASUREMENTS (n = 5). Head length 1.69 ± 0.08 (1.61–1.76); width 1.90 ± 0.12 (1.76–2.10); pronotum length 2.83 ± 0.30 (2.41–3.22); width 5.70 ± 0.46 (5.32–6.45); scutellum length 3.51 ± 0.29 (3.14– 3.95); width 3.33 ± 0.28 (3.06–3.79); length of antennomeres: I 0.30 ± 0.04 (0.26–0.34), II 1.08 ± 0.11 (0.93–1.24), III 1.06 ± 0.10 (0.93–1.16), IV 1.26 ± 0.11 (1.12–1.39), V 1.22 ± 0.09 (1.12–1.31); length of labiomeres: I 0.93 ± 0.12 (0.79–1.12), II 1.06 ± 0.09 (0.94–1.09), III 0.80 ± 0.09 (0.67–0.94), IV 0.83 ± 0.08 (0.71–0.94); length of abdomen 4.67 ± 0.46 (4.03–5.24); width 4.83 ± 0.40 (4.43–5.48); total length 10.03 ± 0.52 (9.35–10.64). GENITALIA. Genital plates cylindrical ( Fig. 10A , C–D, F, I, gp). Parameres long, uniramous, head elongated triangular, dorsally directed, extended beyond pygophore ( Fig. 10 A–H, par). Phallus. Basal theca and thecal shield subequal in length ( Fig. 10 L–R, bt, ts). Vesica subtriangular in dorsal view ( Fig. 10 P–R, v), golf club-shaped in lateral view, with apex obtuse and a central, strongly elevated portion covered by microsculptures ( Fig. 10 L–O, v, m), this central portion broad, rectangular in lateral view, cylindrical in dorsal view; ductus seminis uniform ( Fig. 10 L–R, ds), dorsally directed. Apices of conjunctival lobes globose, endowed with a set of small sculptured processes ( Fig. 10L, N, O , cl, sp). Female MEASUREMENTS (n = 5). Head length 2.12 ± 0.19 (1.83–2.28); width 2.12 ± 0.12 (1.91–2.21); pronotum length 3.17 ± 0.20 (2.82–3.30); width 6.94 ± 0.31 (6.45–7.25); scutellum length 4.43 ± 0.39 (3.79–4.75); width 4.06 ± 0.34 (3.46–4.35); lengths of antennomeres: I 0.38 ± 0.04 (0.34–0.45), II 1.30 ± 0.15 (1.12–1.42), III 1.17 ± 0.14 (0.93–1.31), IV 1.50 ± 0.19 (1.16–1.61), V 1.37 ± 0.15 (1.12–1.50); lengths of labiomeres: I 1.04 ± 0.07 (0.94–1.12), II 1.27 ± 0.15 (1.01–1.38), III 0.93 ± 0.09 (0.78–1.01), IV 0.94 ± 0.11 (0.75–1.05); length of abdomen 6.17 ± 0.34 (5.64–6.45); width 6.27 ± 0.42 (5.64–6.61); total length 11.96 ± 0.83 (10.80–13.06). GENITALIA. Gonocoxites VIII slightly wider than long, posterior margins sinuous ( Fig. 11 A–B, gcVIII). Median and inner duct of vesicular area of uniform diameter ( Fig. 11 C–D, md, id). Fig. 9. Afrius ( Afrius ) purpureus ( Westwood, 1837 ) , variation of the pronotum shape. Scale bars: 2 mm. Fig. 10. Afrius ( Afrius ) purpureus ( Westwood, 1837 ) , male genitalia. A–F . Pygophore in dorsal (A, D), ventral (B, E) and posterior (C, F) views. G–H . Right paramere in lateral views, internal and external respectively. I . Right genital plate, dorso-posterior view. J–L . Phallus in anterior, posterior and lateral views, respectively. M . Detail of the vesica in lateral view, also represented in O. N . Phallus in lateral view. O . Drawing of the phallus. P–R . Phallus in dorsal view. Abbreviations: bf = basal foramen; bp = basal plates; bt = basal theca; cl = conjunctival lobes; db = dorsal border; ds = ductus seminis; er = ejaculatory reservoir; gp = genital plates; il = inferior layer; m = microsculptures; pa = posterolateral angles; par = parameres; ph = phallotheca; sg = secondary gonophore; sp = conjunctival process; ts = thecal shield; v = vesica; vb = ventral border; X = segment X. Scale bars: A–C = 0.5 mm; G–K, N, P–Q = 0.25 mm. Distribution South Africa ( Germar 1838 ), Senegal ( Guérin-Méneville 1844 ), Sierra Leone ( Dallas 1851 ), Guinea ( Stål 1864 ), Equatorial Guinea [as Fernando Po ] ( Stål 1870 ), Ghana ( Reuter 1882 ), Mozambique ( Gerstaecker 1892 ), Zimbabwe [as Mashonaland ] ( Distant 1898 ), Democratic Republic of the Congo ( Distant 1901 ), Equatorial Guinea [as Fernando Po ] ( Stål 1870 ), Ghana ( Reuter 1882 ), Mozambique ( Gerstaecker 1892 ), Benin , Cameroon , Central African Republic [as Haute-Sangha], Eritrea , Ethiopia [as Abyssinia ], Gabon , Ivory Coast [as Grand Bassam], Nigeria [as Benue Niger ], Sierra Leone , South Sudan , Sudan , Tanzania [as Usambara and Kilimanjaro ] (Schouteden 1905a), Kenya [as Leitokitok] ( Schouteden 1910 ), Mali [as Koulikoro] ( Vuillet & Vuillet 1911 ), Guinea-Bissau ( Schouteden 1913b ), Namibia [as Damaraland, Otjiwarongo and Tsumeb] ( Hesse 1925 ), Uganda ( Carpenter 1926 ), Somalia ( Mancini 1937 ), Cape Verde Islands ( Lindberg 1958 or 1959?), Chad ( Couilloud 1989 ), Yemen ( Linnavuori 1989 ), Angola , Botswana , Gambia , Liberia ( Thomas 1994 ), Malawi ( Sileshi et al. 2000 ), Zambia ( Sileshi et al . 2001 ), Republic of Djibouti , Togo ( Maldès & Pluot-Sigwalt 2004 ), Republic of the Congo [as Haut Oubangui?] ( Maldès & Pluot-Sigwalt 2004 ; also in Robertson 2009 as “ Congo Brazz.”), Niger ( Robertson 2009 ; in error?), Canary Islands (new record) ( Table 1 , Fig. 1 ). Fig.11. Afrius ( Afrius ) purpureus ( Westwood, 1837 ) ,female genitalia. A–B .Genital plates,ventroposterior view. C–D . Inner genitalia, ventral view. Abbreviations: cs = capsula seminalis; gcVIII = gonocoxites VIII; gcIX = gonocoxites IX; gpIX = gonapophyses IX; id = inner duct; laVIII = laterotergites VIII; laIX = laterotergites IX; md = median duct of vesicular area; od = outer duct of vesicular area; pi = pars intermedialis; tgIX = secondary thickening of gonapophyses IX; vi = thickening of vaginal intima; X = segment X. Scale bars: A–B = 1 mm; C–D = 0.5 mm. Remarks Intraspecific variability in the colour and general morphology has been demonstrated for A. (Afrius) purpureus (e.g., Linnavuori 1989 ; Schouteden 1905a; Van Heerden 1931 ; Villiers 1952b ) as well as for other pentatomids, such as Nezara viridula (Linnaeus, 1758) (e.g., Freeman 1940 ; Kiritani 1970 ; Ohno & Alam 1992 ; Vivan & Panizzi 2002 ; More et al. 2017 ) and Stiretrus decemguttatus (Lepeletier & Serville, 1828) ( Paleari 2013 ) for colour pattern, and Pinthaeus sanguinipes (Fabricius, 1781) for variation in the pronotum, reported by Zhao et al. (2013) . Besides, we could not find any connection between the polymorphism and the geographic distribution of A. ( Afrius ) purpureus , i.e., specimens that we have examined, despite different colour patterns and pronotum shapes, are sympatric. The localities where they were collected are denoted by the green dots on Fig. 1 . There has been considerable difference of opinion regarding the name used for this species, i.e., either yolofa /- us , purpurea /- us or figurata /- us and, sometimes, the latter used as a variety or subspecies of the second one (see review of the taxonomic history of the species above). Dallas (1851) was the first to synonymize figuratus and yolofa ; he stated the date of publication of Pentatoma yolofa as 1830, while Stål (1864) stated it as 1829. Subsequent authors (including Stål 1870 ) accepted 1838 as the year of publication for yolofa and therefore they recognized either figuratus or purpureus as the valid name, Wallengren (1875) may have been the last author to use yolofa (misspelled as Ylofa ) as a valid name while Larousse (1890) gave a brief description of it under the genus Canthecona (strangely, this entry first mentions that the type species of Canthecona is from Senegal and thereafter describes C . yolofa , as though implying it were the type species; the type species of Canthecona Amyot & Serville, 1843 is, however, C . discolor (Palisot de Beauvois, 1811) , described from the Kingdom of Oware, now Southwest Nigeria ). Dupuis (1952) demonstrated the priority of Pentatoma yolofa Guérin-Ménéville, 1831 over Pentatoma purpurea Westwood, 1837 and Asopus figuratus Germar, 1838 . Thomas (1994) adopted the combination suggested by Dupuis (1952) , Afrius yolofus . This was, as well, used by subsequent authors ( Maldès & Pluot-Sigwalt 2004 ; Rebagliati et al. 2005 ; Kerisew 2011 ). Some others ( Sileshi et al. 2001 , 2004 ; Kerzhner et al. 2004 ; Rider 2006 ; Robertson 2009 ; Matesco et al. 2014 ), however, chose the combination Afrius yolofa , while others again still used purpureus and/or figuratus after 1952 (e.g., Mancini 1953 ; Villiers 1954 , 1956 ; Leston 1954 ; Lindberg 1958 ; Schouteden 1963 , 1964 ; Gillon 1972 ; Linnavuori 1975 , 1976 , 1982 ; Bourdouxhe & Jolivet 1981 ; Nuamah 1982 ; Matanmi & Hassan 1987 ; Balsbaugh 1988 ; Couilloud 1989 ; Quicke et al. 1992 ; Schaefer 1996 ; Tchibozo & Braet 2004 ; Poutouli et al. 2011 ; Agboton et al. 2014 ; Crétenet & Gourlot 2015 ) and yet others treated figuratus as a variety or a subspecies of purpureus (Schouteden 1905a, 1907a; Kirkaldy 1909 ; Schouteden 1909 ; Mancini 1937 ; Le Pelley 1959 ; Linnavuori 1989 ). Others still have used two or three combinations in the same work, possibly because they were reporting facts from primary sources, using the names as they were in the sources, and were not aware that the species were the same ( Le Pelley 1959 ; Schaefer 1996 ). The different combinations used for one and the same species clearly are the results of a few misconceptions: when and how yolofa , figuratus and purpureus were synonymized, the problematic dating of yolofa and figuratus and the status of yolofa as an adjective or a noun in apposition.