Invreiella satrapa ( Gerstaecker, 1874 )

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, Zootaxa 4894 (2), pp. 151-205 : 191-193

11. Invreiella satrapa ( Gerstaecker, 1874)

( Figs 5–8 , 19 , 29, 31 , 63 , 75 , 91 , 106 , 122 , 136 ; Map 6)

Mutilla satrapa Gerstaecker, 1874: 65 [lectotype ♀ (ZMHB)]. Dalla Torre 1897: 83.

- Sphaerophthalma [sic] satrapa: Cameron 1895: 364 .

- Ephuta (Ephuta) satrapa: André 1902: 63 .

- Pseudomethoca satrapa: Mickel 1937: 181 ; Mickel 1964: 168.

- Invreiella satrapa: Suárez 1966: 475 ; Nonveiller 1990: 41; Pagliano 2005: 272; Lelej & Brothers 2008: 29; Quintero & Cambra 2011: 219; Cambra et al. 2014: 296; Pagliano et al. 2020: 102.

Mutilla macrocephala Smith, 1879: 223 [holotype ♀ (NHMUK)]. Junior subjective synonym of Pseudomethoca satrapa ( Gerstaecker, 1874) according to Mickel 1964: 168. Synonym status confirmed.

- Sphaerophthalma [sic] macrocephala: Blake 1886: 190 , 256.

- Ephuta (Ephuta) macrocephala: André 1902: 61 .

-? Pseudomethoca macrocephala: Mickel 1924: 46 .

Mutilla ganahlii Dalla Torre, 1897: 42 [new name for Mutilla macrocephala Smith, 1879 , nec Olivier, 1811]. Junior subjective synonym of Pseudomethoca satrapa ( Gerstaecker, 1874) according to Mickel 1964: 168. Synonym status confirmed.

Diagnosis (female). This species is distinguished from other members of the I. satrapa species-group by the following combination of characters: clypeus elongate, extending well-beyond base of mandibular socket (i.e. fig. 24; fig. 91), with medioapical tubercles basally conjoined and dorsoventrally triangular in shape, genal process spinose and straight (figs 29, 75), pronotal carina strongly lobate posteriorly and projecting away from mesosoma, overlapping and obscuring dorsal portion of pronotal-mesopleural suture (figs 31, 106, 122), and extreme apical margin of T2 with dark orange integument (figs 19, 136).

Redescription (female). Body length 8.26–12.95 mm.

Head: Head 1.03–1.19 × as wide as mesosoma. Vertex and frons contiguously punctate. Antennal scrobe carina straight to weakly arcuate, slightly more prominent internally, with inner tip of carina well-separated from antennal rim. Frons ranging from weakly to moderately transversely recessed and concave below antennal scrobe carina, antennal rim consequently slightly recessed and slightly visible to recessed and not visible when head viewed laterally. Antennal rim apically glabrous, rounded and not tuberculate. F1 1.67–2.13 × as long as F2. Clypeus plate-like and medially flattened, coarsely rugose-granulate, with central cluster of long raised setae, apical margin medially elongate and coming to distinct apex, with two close medioapical tubercles that are anterad-projecting, dorsoventrally triangular in shape, and basally conjoined. Mandible acuminate. Distance from posterior margin of eye to posterolateral corner of head 1.37–1.9 × maximum diameter of eye. Genal process spinose and straight, posterior genal carina nearly straight. Gena with separate yet close moderate punctures laterally, interpunctal space smooth. Postgena transversely rugose-striate.

Mesosoma: Mesosoma 1.2–1.42 × as wide as long. Dorsum and posterior face of mesosoma densely, contiguously punctate, surfaces coarse. Anterodorsolateral margin of pronotum outcurved. Pronotal carina prominent, glabrous, visible both dorsally and laterally, strongly lobate posteriorly and projecting away from mesosoma, overlapping and obscuring dorsal portion of pronotal-mesopleural suture. Tubercle anterior of propodeal spiracle present. Width of propodeum greater than distance between propodeal spiracles in dorsal view. Lateral face of pronotum rugose-striate, with scattered moderate punctures, lateral face posteriorly carinate along pronotal-mesopleural suture in addition to aforementioned lobate structure of pronotal carina. Mesopleuron microgranulate to granulate anteriorly, with few punctures, posteriorly striate-rugose and weakly glabrous. Mesopleuron with vertical column of punctures weakly tuberculate, not forming anterior or posterior carinae. Mesopleuron posteriorly carinate along ventral one-half to two-thirds of mesopleural-metapleural suture, dorsal portion of suture obscure, faintly present. Metapleuron with ventral one-third to one-half striate-rugose with scattered punctures, dorsal one-third to one-half microgranulate and shining. Metapleural-propodeal suture with striae perpendicularly overlapping suture along ventral one-third to one-half. Lateral face of propodeum posteriorly with ventral one-third to one-half striate-rugose with scattered punctures, anteriorly with dorsal one-third to one-half microgranulate and shining. Coxae coarsely sculptured, with small to large, deep punctures.

Metasoma: T1–5 densely, contiguously punctate. Visible portion of pygidial plate not obscured by setal hood transversely rugose to rugose-granulate. S2 densely punctate, punctures near contiguous in some areas, basolateral concave area less densely 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 orange to black: antenna, antennal rim, medioapical tubercles of clypeus, apical one-third to near entirety of mandible, legs, T1, extreme apical margins of T2 and S2, T3–6, and apical margins of S3–6; pleura, coxae, and S1 vary between orange and dark red-brown; posterior and lateral faces of propodeum vary between orange and nearly black.

Pubescence: Frons ventrally with pale to light orange setae. Frons dorsally and vertex (including posterolateral corner of head) covered with mostly decumbent orange-red setae, with few scattered fuscous setae. Clypeus with long pale or fuscous setae medially. Mandible with dorsal and ventral longitudinal row of light orange setae. Remainder of head with whitish setae. Dorsum of mesosoma mostly covered with decumbent orange-red setae, dorsal edges (except pronotum) with sparse raised fuscous setae, particularly at top of posterior face of propodeum. Pronotum without black setae, rather with orange setae. Remainder of mesosoma including legs with whitish setae (except tarsi with dense orange bristles). Anterior face of T1 with whitish setae. T1 apically fringed with wide band of black setae medially, laterally fringed with whitish setae. Dorsum of T2 covered with orange-red setae. Felt line of T2 and surrounding lateral area with whitish setae. T2 apically fringed with moderately wide band of black setae medially, not coming to median point, laterally fringed with whitish setae. T3–5 mostly covered and apically fringed with whitish setae, with few scattered dark raised setae and apical fringe with median patch of black setae, these black setal patches mostly consistent in width between T3–5, with T5 black setal patch usually covering median section of sclerite. T6 with pale yellow-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 Greek satrápçs, meaning governor of a province.

Distribution: Mexico (Coahuila, Durango, Guanajuato, Hidalgo, Mexico, Michoacan, San Luis Potosi, 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).

Host(s): Unknown.

Remarks: Gerstaecker (1874) based his description of M. satrapa on at least two specimens, as he provided a length range of “11–14 mill.” Upon examining the type of M. satrapa , it was observed that C. Mickel designated it as a lectotype, which he never explicitly published. Two additional syntypes were discovered at ZMHB (L. Kirschey, pers. comm.). In Mickel (1964), implicit reference is made to the location of the type of M. satrapa , and the word ‘type’ is missing from the treatment of this species. According to the ICZN (1999), article 74.5: Lectotype designations before 2000: “In a lectotype designation made before 2000, either the term “ lectotype,” or an exact translation or equivalent expression (e.g. “the type”), must have been used or the author must have unambiguously selected a particular syntype to act as the unique name-bearing type of the taxon.” Consequently, Mickel’s unpublished lectotype designation for M. satrapa is here validated. There is some natural wear to the lectotype of I. satrapa : the apices of the mandibles and medioapical tubercles of the clypeus are worn down. This gives the impression that the clypeal tubercles are dorsoventrally semi-circular in shape, rather than triangular, which is not the case for this species. Additionally, the apices of both genal processes are broken off. Among the specimens of I. satrapa that have been examined, minor asymmetry in the shape of the genal carinae and processes has been observed to varying degrees.

The holotype specimen of Mutilla macrocephala Smith, 1879 and its collection data are noteworthy and are here expanded upon. Smith (1879) noted the distribution of this species as “Orizaba,” a city in Veracruz, Mexico; interestingly, the only locality included on the holotype’s label data is “Mex.” Orizaba is a common locality among Hymenoptera that F. Smith had described and we maintain this locality here despite the type not explicity being labeled as such. Additionally, the holotype of M. macrocephala is unusual in that its frons is more transversely recessed and concave below the scrobe carinae than most other specimens of I. satrapa .

Lastly, there are two records of this species being collected on plants. One female was collected on Bidens pilosa Linnaeus (beggar-ticks/Spanish needle) and another on a pepper tree (both SEMC).

Material examined ( I. satrapa , 28♀).

Lectotype ( Mutilla satrapa Gerstaecker ) (designated by C.E. Mickel (unpublished), here designated) (examined, figs 5–8): ♀ ( ZMHB) [apices of both genal processes broken off; entire right proleg missing], [label 1 (green):] Mexico / Ehrenberg S. [// label 2 (tan):] 6585 [// label 3 (red):] Type [// label 4 (red):] Lectotypus / C.E. Mickel [// label 5 (tan):] Zool. Mus. / Berlin [// label 6 (white):] Mutilla satrapa . [( MEXICO: Unknown : Ehrenberg S. (1♀ – ZMHB))]

Syntypes ( Mutilla satrapa Gerstaecker ) (examined via photographs, consequently not designated as paralectotypes): MEXICO: Unknown: Ehrenberg S. (2♀ – ZMHB) .

Holotype ( Mutilla macrocephala Smith ) (examined): ♀ ( NHMUK) [F2–10 of left antenna missing; entire right antenna missing; most of femur, entire tibia, and tarsus of left proleg missing; tarsus missing of left mesoleg missing; tarsomere #5 of left metaleg missing; tarsus of right proleg missing; tarsomeres #4–5 of right mesoleg missing; tarsomere #5 of right metaleg missing], [label 1 (circular, light blue, obverse):] Mex [label 1 (reverse):] 65 / 31 [// label 2 (tan):] Mutilla / macrocephala / (Type) Sm. [// label 3 (circular, tan center with red outline):] Type [// label 4 (tan, folded, inner side):] Smith’s description of / macrocephala does / not correspond with / the type specimen / The latter is con- / -specific with jocularis, Cameron / C.L.B. [label 4 (outer side):] This is not / correct. / C.E. Mickel / 1930 [// label 5 (tan):] B.M. TYPE / HYM. / 15.1043. [( MEXICO: Veracruz: [Orizaba], [18.850°N 97.103°W] (1♀ – NHMUK))]

Additional specimens (non-types, examined or referenced*): MEXICO: Coahuila: Saltillo , [25.430°N 100.977°W], Gribodo, MNCN_ Ent 171986 (1♀ – MNCN *) . Durango: Victoria de Durango, 31.7 mi. N, [24.403°N 104.696°W], 22.Jul.1982, A.J. Gilbert (1♀ –0006578– CSCA) . Guanajuato: Roque , [20.581°N 100.838°W], 31.Oct.1964, H. Velasco, “HOSP. plantas silvestres” (1♀ –0000045– EMUS) ; San José de Allende, [20.856°N 101.033°W], oak forest, 9000 ft., 10.Aug.1988, G.B. Edwards (1♀ –0000047– FSCA) . Hidalgo: Atotonilco El Grande, Sanctórum , [20.301°N 98.777°W], 12.Sep.1992, L. Godinez, “#735, ex. Bidens pilosa L.” (1♀ – SEMC) ; Pachuca, [20.104°N 98.763°W], 7900 ft., 28.Jul.1954, University of Kansas Mexico Expedition (1♀ – DJBC *) ; Pa-chuca, 4 mi. W, [20.128°N 98.838°W], 7900 ft., 16.Jun.1961, University of Kansas Mexico Expedition (1♀ – DJBC *) . México: Mexico City, [19.432°N 99.133°W], H. Höge (1♀ –0000040– UMSP; 1♀ –0000058– NHMUK) ; Mexico City, Mixcoac , [19.372°N 99.190°W], H.F. Wickham (1♀ –0000041– UMSP) ; Tepexpan , [19.614°N 98.936°W], 26.Jul.1963, F.D. Parker & L.A. Stange (1♀ –0000046– EMUS; 1♀ –0006577– UCDC) ; Venta de Carpio, 0.6 mi. SE, [19.615°N 98.999°W], 7800 ft., 06.Aug.1972, R.R. & M.E. Murray , “ME-68” (1♀ –0000043– TAMU; 1♀ – 0000044– GCWC) ; Zoquiapan [National Park], [19.383°N 98.708°W], 7000 ft., 14.Jul.1953, University of Kansas Mexico Expedition , “taken on pepper tree” (1♀ – SEMC) . Michoacán: Tzintzunzán , 1 km S, [19.616°N 101.576°W], 03.Nov.1992, F.A. Noguera & A. Rodríguez (1♀ –0000042– EBCC) . San Luis Potosí: [San Luis Potosí], [22.156°N 100.985°W], E. Palmer (2♀ –0000059, 0007210– NHMUK) ; San Luis Potosí, 20 mi. SW, [22.009°N 101.247°W], 6800 ft., 25.Jul.1962, University of Kansas Mexico Expedition (1♀ – DJBC *; 1♀ – SEMC) ; San Luis Potosí, 29 mi. SW, [21.929°N 101.358°W], 6800 ft., 25.Jul.1962, University of Kansas Mexico Expedition (1♀ – DJBC *) . Unknown: (1♀ –0000050– NHMD) . Veracruz: Perote, 10 mi. SW, [19.484°N 97.364°W], 27.Jul.1974, Clark, Murray , Ashe , & Schaffner (1♀ –0000049– DGMC) . Zacatecas: Fresnillo , 9 mi. S, [23.040°N 102.874°W], 20.Aug.1956, D.D. Linsdale, EMEC 1157454 (1♀ –0000048– EMEC) .

Figures

FIGURES 1–8. Invreiella cardinalis, holotype female: 1–dorsal habitus, 2–lateral habitus, 3–clypeus, 4–labels. I. satrapa, lectotype female: 5–dorsal habitus, 6–lateral habitus, 7–clypeus, 8–labels. Scale bars = 2 mm.

FIGURES 9–23. Female habitus, dorsal view. Scale bars = 2 mm.

FIGURES 24–31. Female clypeus: 24–I. chihuahuensis (0000031), 25–I. breviclypeata (0000003), 26–I. cardinalis (0000018), 27–I. australis (0000001). Female head, left lateral view: 28–I. chihuahuensis (0000031), 29–I. satrapa (0000049). Female pronotum, left lateral view: 30–I. chihuahuensis (0000031), 31–I. satrapa (0000049).

FIGURES 55–63. Female head, left lateral view: 55–I. cephalargia (0000024), 56–I. tequila (0000053). Female mesopleuron, left lateral view: 57–I. suarezi (0000051), 58–I. tequila (0000053). Female mandible, anterior view: 59–I. tequila (0000053), 60–I. suarezi (0000051). Female mesosoma, left lateral view: 61–I. manleyi (0000037), 62–I. erythrocephala (0000034). Female procoxae, anterior view: 63–I. satrapa (0000045).

FIGURES 64–79. Invreiella australis (0000001): 64–female habitus, lateral view. Female head, left lateral view: 65–I. australis (0000001), 66–I. bimaculata (0000002), 67–I. mesomexicana (0000035), 68–I. acuminata (0000056), 69–I. cardinalis (0000018), 70–I. cuernavaca (0000032), 71–I. cephalargia (0000024), 72–I. manleyi (0000037), 73–I. breviclypeata (0000003), 74–I. chihuahuensis (0000031), 75–I. satrapa (0000049), 76–I. erythrocephala (0000034), 77–I. suarezi (0000051), 78–I. tequila (0000053), 79–Pseudomethoca frigida.

FIGURES 80–94. Invreiella australis (0000001): 80–female head, anterior view. Femaleclypeus:81–I.australis (0000001), 82– I. bimaculata (0000002), 83–I. mesomexicana (0000035), 84–I. acuminata (0000056), 85–I. cardinalis (0000018), 86–I. cuernavaca (0000032), 87–I. cephalargia (0000024), 88–I. manleyi (0000037), 89–I. breviclypeata (0000003), 90–I. chihuahuensis (0000031), 91–I. satrapa (0000049), 92–I. erythrocephala (0000034), 93–I. suarezi (0000051), 94–I. tequila (0000053).

FIGURES 95–110. Invreiella australis (0000001): 95–female habitus, dorsal view. Female mesosoma, left dorsal view: 96–I. australis (0000001), 97–I. bimaculata (0000002), 98–I. mesomexicana (0000035), 99–I. acuminata (0000056), 100–I. cardinalis (0000018), 101–I. cuernavaca (0000032), 102–I. cephalargia (0000024), 103–I. manleyi (0000037), 104–I. breviclypeata (000003), 105–I. chihuahuensis (0000031), 106–I. satrapa (0000049), 107–I. erythrocephala (0000034), 108–I. suarezi (0000051), 109–I. tequila (0000053), 110–Pseudomethoca frigida.

FIGURES 111–125. Invreiella australis (0000001): 111–female habitus, lateral view. Female mesosoma, left lateral view: 112–I. australis (0000001), 113–I. bimaculata (0000002), 114–I. mesomexicana (0000035), 115–I. acuminata (0000056), 116–I. cardinalis (0000018), 117–I. cuernavaca (0000032), 118–I. cephalargia (0000024), 119–I. manleyi (0000037), 120–I. breviclypeata (0000003), 121–I. chihuahuensis (0000031), 122–I. satrapa (0000049), 123–I. erythrocephala (0000034), 124–I. suarezi (0000051), 125–I. tequila (0000053).

FIGURES 134–136. Female habitus, dorsal view: Invreiella satrapa species-group. Scale bars = 2 mm. MAP 6. Geographic distribution of the Invreiella satrapa species-group.