Invreiella cardinalis

5. Invreiella cardinalis (Gerstaecker, 1874) (Figs 1–4, 13, 26, 33, 36, 37, 39, 69, 85, 100, 116, 130; Map 4) Mutilla cardinalis Gerstaecker, 1874: 64 [holotype ♀ (ZMHB)]. Dalla Torre 1897: 21. - Sphaerophthalma [sic] cardinalis: Cameron 1895: 364. - Ephuta (Ephuta) cardinalis: André 1902: 58. - Pseudomethoca cardinalis: Mickel 1964: 166. - Invreiella cardinalis: Suárez 1966: 475; Nonveiller 1990: 41; Quintero & Cambra 2011: 218; Pagliano et al. 2020: 102. Sphaerophthalma [sic] jocularis Cameron, 1894: 315; Tab. 14, fig. 3 [holotype ♀ (NHMUK)]. Junior subjective synonym of Pseudomethoca cardinalis (Gerstaecker, 1874) according to Mickel 1964: 166. Synonym status confirmed. - Ephuta (Ephuta) jocularis: André 1902: 61. -? Pseudomethoca jocularis: Mickel 1924: 46. - Pseudomethoca jocularis: Suárez 1962: 120. Invreiella curoei Quintero & Cambra, 2011: 218 [holotype ♀ (MIUP)]. Pagliano et al. 2020: 102. New synonym. Diagnosis (female). This species is distinguished from other members of the I. cardinalis species-group by the following combination of characters: antennal scrobe carina straight (figs 39, 85), antennal rim weakly tuberculate, genal process spinose and curved with posterior genal carina sinuate (figs 33, 69), with carina anterior to process crenulate-tuberculate, anterodorsolateral margin of pronotum outcurved (figs 36, 100), dorsolateral margin of mesosoma, from pronotal carina to propodeal spiracle, distinctly sinuate and undulating in outline (figs 36, 100), width of propodeum greater than distance between propodeal spiracles in dorsal view (figs 36, 100), mesopleuron with ventral one-third coarsely sculptured, rugose-granulate with weak rugae and few moderate punctures (figs 37, 116), and integument of dorsum of head, mesosoma, and center of T2 orange-red (figs 13, 130). Redescription (female). Body length 8.51–11.94 mm. Head: Head 1.16–1.31 × as wide as mesosoma. Vertex and frons contiguously punctate. Antennal scrobe carina straight, with inner tip of carina separated from antennal rim and slightly more prominent than rest of carina. Frons weakly transversely recessed and concave below antennal scrobe carina, antennal rim consequently not recessed and is visible when head viewed laterally. Antennal rim apically glabrous, weakly tuberculate. F1 1.83–2.89 × as long as F2. Clypeus concave, with transverse arcuate carina broadly interrupted medially, area merely punctate, resulting in bilateral carinate processes each with enlarged tubercle ventrad to process. Mandible acuminate. Distance from posterior margin of eye to posterolateral corner of head 1.43–2.05 × maximum diameter of eye. Genal process spinose and curved, posterior genal carina sinuate, carina anterior to process crenulate-tuberculate. Gena densely punctate laterally, punctures large and nearly contiguous, interpunctal space smooth; Postgena transversely rugose-striate medially. Mesosoma: Mesosoma 1.10–1.28 × as wide as long. Dorsum and posterior face of mesosoma densely, contiguously punctate, surfaces coarse. Anterodorsolateral margin of pronotum outcurved. Pronotal carina prominent, glabrous, weakly crenulate, visible both dorsally and laterally, not strongly protruding posteriorly in dorsal view and not entirely obscuring apical portion of pronotal-mesopleural suture. Dorsolateral margin of mesosoma, from pronotal carina to propodeal spiracle, distinctly sinuate and undulating in outline in dorsal view; tubercle anterior of propodeal spiracle obscure. Width of propodeum greater than distance between propodeal spiracles in dorsal view. Lateral face of pronotum weakly to moderately rugose-striate, with moderate punctures between rugae, lateral face posteriorly carinate along pronotal-mesopleural suture. Mesopleuron with dorsal two-thirds microgranulate absent. Metapleuron rugose-striate dorsad metacoxa, with few micro- and moderate punctures, remainder of sclerite weakly microgranulate, near glabrous to glabrous. Metapleural-propodeal suture with overlapping striae at ventral one-third. Lateral face of propodeum posteriorly and with ventral one-third to one-half rugose-striate, with moderate punctures between rugae, dorsal one-third to one-half weakly microgranulate, near glabrous to glabrous. Coxae coarsely sculptured, with small to moderate deep punctures. Metasoma: T1–5 densely, contiguously punctate. Visible portion of pygidial plate not obscured by setal hood rugose-granulate. S2 densely punctate, punctures near contiguous in some areas, basolateral concave area sparsely punctate, nearly glabrous between punctures. S3–6 densely punctate, coarsely microgranulate between punctures. Hypopygium with slightly arcuate transverse row of setae near apical margin, lateral setae longer than median setae. Integument coloration: Orange, except the following ranging from dark red-brown to black: antenna, antennal rim, apex and base of mandible, tip of genal process, legs including coxae and trochanters (rarely femora orange), and metasoma (excluding two conjoined orange maculae on T2 and rarely T1 orange). Pubescence: Frons ventrally with whitish setae, medially with few fuscous setae. Frons dorsally and vertex mostly covered with decumbent, and some raised, light yellow to orange setae. Posterolateral corner of head with few scattered raised fuscous setae. Mandible with dorsal and ventral longitudinal row of light orange setae. Remainder of head with whitish setae. Dorsum of mesosoma mostly covered with short decumbent light yellow to orange-red setae, dorsolateral edges with raised fuscous to whitish setae. Pronotum with dorsal transverse band of black setae, or only orange setae, with area between epaulet and pronotal spiracle with few whitish setae. Remainder of mesosoma including legs with whitish setae (except tarsi with orange bristles). Anterior face of T1 with whitish setae. T1 apically fringed with wide median band of black setae, laterally fringed with whitish setae. T2 with decumbent light yellow to orange setae overlapping conjoined orange integumental maculae, with maculae anteriorly and posteriorly surrounded with black setae. Felt line of T2 and surrounding lateral area with whitish setae. T2 apically fringed with wide band of black setae medially, laterally fringed with whitish setae. T3 covered and fringed with whitish setae, few scattered raised fuscous setae present. T4 similar to T3 except apically fringed with small patch of black setae medially. T5 mostly covered and fringed with black setae medially, laterally fringed with whitish setae. T6 with light orange setae surrounding pygidial plate, partially obscuring basal half. Remainder of metasoma with whitish setae. MALE. Unknown. Etymology: The specific epithet is likely derived from the Latin cardinalis in reference to the red-orange coloration of this species. Distribution: Mexico (Durango, Guanajuato, Guerrero, Jalisco, Mexico, Morelos, Oaxaca, Puebla, Veracruz, and Zacatecas). Biogeography: Nearctic region (Chihuahuan Desert province); Mexican transition zone (Sierra Madre del Sur province, Sierra Madre Occidental province, Sierra Madre Oriental province, Transmexican Volcanic Belt province); Neotropical region (Balsas Basin province). Host(s): Unknown. Remarks: In contrast to I. satrapa, Gerstaecker described I. cardinalis based on a single specimen. There are no syntypes of I. cardinalis at ZMHB (Lukas Kirschey, pers. comm.). Further, Gerstaecker gives a single length as part of the description, rather than a length range as in I. satrapa, implying that a single specimen was examined by him. There is some variation in setal coloration of the dorsum of the pronotum, which can range from light yellow to orange to black. One small female from Veracruz, Mexico (0000016– CASC) has the genal carina less strongly recurved and the genal process slightly reduced, unlike those of the other specimens examined. Another specimen from Durango, Mexico (0007207– NHMUK) also has the genal carina less recurved than most specimens, with there being minor asymmetry in curvature (the left carina slightly less recurved than the right carina). Further, this specimen has the integument of the femora and T1 orange, which is unusual for this species. Label data on one specimen (0000014–EMEC) indicates that it was collected on flowers of the Mexican kidneywood tree, Eysenhardtia polystachya (Ortega) Sarg.; this female was likely nectaring. Another specimen was collected on an Ipomoea sp. (morning glory) (SEMC). Material examined (I. cardinalis, 31♀). Holotype #1 (Mutilla cardinalis Gerstaecker) (examined, figs 1–4): ♀ (ZMHB) [head and propleura (with attached prolegs) glued onto specimen], [label 1 (green):] Mexico. / Jalappa / Deppe. [// label 2 (red):] Type [// label 3 (tan):] 6584 [// label 4 (tan):] Zool. Mus. / Berlin [// label 5 (white):] Mutilla cardinalis. [(MEXICO: Veracruz: Xalapa, [19.543°N 96.910°W], F. Deppe (1♀ – ZMHB))] Holotype #2 (Sphaerophthalma [sic] jocularis Cameron) (examined): ♀ (NHMUK) [F7–10 of right antenna missing; both prolegs entirely missing (apparently glued back onto specimen at one point); mesosoma dorsally cracked due to pin; propodeum mostly covered with thick layer of dirt], [label 1 (tan):] Amula, / Guerrero, / 6000 ft. / Sept. H. H. Smith. [// label 2 (tan):] B.C.A.Hymen.II. // Sphaerophthalma / jocularis, / Cam. [// label 3 (tan, folded):] Sphaerophthalma / jocularis / Cam. type / BCA ii 315 [// label 4 (tan):] 1903–297. [// label 5 (circular, tan center with red outline):] Type [// label 6 (tan):] B.M. TYPE / HYM. / 15.1042. [(MEXICO: Guerrero: Amula village, [17.633°N 99.250°W], 1828.8 m, Sep., H.H. Smith (1♀ – NHMUK))] Holotype #3 (Invreiella curoei Quintero & Cambra) (examined): ♀ (MIUP). [(MEXICO: Puebla: Tehuacán, [18.466°N 97.400°W], 5500 ft., 06.Jul.1941, H. Seevers (1♀ – MIUP))] Additional specimens (non-types, examined or referenced*): MEXICO: Durango: Ciudad Lerdo, [25.544°N 103.526°W], H. Höge (1♀ –0007207– NHMUK). Guanajuato: [Guanajuato], [21.019°N 101.257°W], E. Dugès (1♀ –0006575– RBINS). Guerrero: Chichihualco, 18 mi. W, [17.587°N 99.799°W], 1970 m, 28.Aug.1977, E.I. Schlinger, EMEC 1135899 (1♀ –0000015– EMEC); Tixtla, 10 mi. E, [17.569°N 99.247°W], 16.Jul.1983, J. Chem-sak, A. & M. Michelbacher, EMEC 1135898 (1♀ –0000013– EMEC). Jalisco: Magdalena, [20.910°N 103.969°W], 15.Jul.1965, W.F. Chamberlain (1♀ –0000018– DGMC); Teocaltiche, Rancho La Quinta, [21.433°N 102.571°W], 5600 ft., 25.Aug.1979, B. Villegas (1♀ –0006576– UCDC); Tepatitlán de Morelos, [20.809°N 102.765°W], 2011.7 m, 20.Aug.1954, J.G. Chillcott (1♀ –0000007– CNC); Tuxpan, Atenquique, [19.529°N 103.442°W], H. Höge (1♀ –0000061– NHMUK). México: Temascaltepec, [19.043°N 100.041°W], 1931, G.D. Hinton (2♀ –0000009, 0000010– EMUS). Morelos: Cuernavaca, [18.924°N 99.221°W], 08.Nov.–06.Dec.1987, F.D. Parker (1♀ –0000008– EMUS). Oaxaca: Monte Albán, [17.043°N 96.767°W], 1900 m, 23.Dec.1990, L. Godinez, #652, “ex. Ipomoea sp.” (1♀ – SEMC). Puebla: Tehuacán, [18.466°N 97.400°W], 12.Jul.1935, A.E. Pritchard (1♀ –0000005– UMSP); 23.Jun.1951, P.D. Hurd, “on flowers of Eysenhardtia polystachya (Ort.),” EMEC 1135783 (1♀ –0000014– EMEC). Unknown: (2♀ –0000004, 0000006– UMSP); (1♀ –0000011– NHMD); (1♀ –0000012– NMNH); (3♀ –0000060, 0007208, 0007209– NHMUK); Sallé, MNCN_Ent 171985 (1♀ – MNCN *). Veracruz: Citlaltépetl, Pico de Orizaba, plateau, [19.013°N 97.266°W], 1524 m, 03.Aug.1964, L.W. Swan (1♀ –0000016– CASC). Zacatecas: Moyahua de Estrada, [21.265°N 103.165°W], 07.Aug.1965, A.R. Gillogly (3♀ – DJBC *; 1♀ – SEMC); Nochistlán, [21.363°N 102.843°W], 23.Aug.1970, B.L. Villegas (1♀ –0000017– DGMC).Published as part of Waldren, George C., Williams, Kevin A., Cambra, Roberto A. & Pitts, James P., 2020, Systematic revision of the North American velvet ant genus Invreiella Suárez (Hymenoptera: Mutillidae) with description of eleven new species, pp. 151-205 in Zootaxa 4894 (2) on pages 177-180, DOI: 10.11646/zootaxa.4894.2.1, http://zenodo.org/record/431561