Smicromyrme (Astomyrme) pusillus

Matias, Rafael, 2024, The subgenus Astomyrme Schwartz, 1984 in the Iberian Peninsula: associating Physetopoda pusilla (Klug in Waltl, 1835) with Smicromyrme suberratus Invrea, 1957, and description of the hitherto unknown female of S. matritentis (Mercet, 1905) (Hymenoptera: Mutillidae), Zootaxa 5480 (1), pp. 1-62 : 17-33

Smicromyrme (Astomyrme) pusillus (Klug in Waltl, 1835), comb. nov.

( Figs 4–10 , 19A–B , 21A–D )

Mutilla pusilla Klug in Waltl, 1835: 88, ♀ (original combination), type locality: Puerto Real (Cádiz, Spain) [vii.1829, J. Waltl leg.], holotype in Museum für Naturkunde Berlin ( Germany). Rosenhauer 1856: 372 (♀, commented); Morawitz 1865: 132 (♀); André 1899–1902: 307 (♀?; probable misidentifications according to Bischoff 1933 and Invrea 1960); Mercet 1905: 491 (♀, in part?, discussed, listed); Berland 1925: 330 (♀ in part?, nec ♂);? Giner Marí 1942: 72 (♀, in part?, listed); Ceballos 1959: 223 (in part?, catalogued).

Mutilla usilla [sic] Klug in Waltl, 1835: 92, ♀ (original description); Klug in Silbermann 1837: 159 (♀).

Mutilla subcomata Radoszkowski, 1888: 347 (♂?, nec ♀); synonymized with Smicromyrme suberrata by Invrea 1957: 198. André 1899–1902: 320 (♂, nec ♀; in part?); Mercet 1905: 492 (♂?, nec ♀). New synonymy.

Mutilla quadrimaculata : André 1896: 271 (discussed; probable misidentification).

Mutilla pussilla [sic]: Mercet 1902: 311 (mentioned);? Antiga y Bofill 1904: 10 (♀).

Smicromyrme pusilla : Bischoff 1930: 2 (♀, listed); Bischoff 1933: 73 (♀, nec ♂; discussed); Giner Marí 1944: 85 (♀, in part?, nec ♂; discussed); Suárez 1952: 82 (♀, in part?, commented); Ceballos 1959: 223 (catalogued); Mingo & Compte 1963: 87 (♀, discussed).

Smicromyrme subcomata : Giner Marí 1944: 92 (♂, nec ♀); Invrea 1950: 27 (♂); Diniz 1989: 30 (♂, probably this species); Suárez 1952: 83 (♂ in part?, nec ♀).

Smicromyrme suberrata Invrea, 1957: 199 , ♂, type locality: Ghedahia (west Sirtica, Libya), 2.v.1939, Krüger leg., holotype in Museo Civico di Storia Naturale “G. Doria” (Genova, Italy); replacement name for Mutilla subcomata ♂ sensu Radoszkowski 1888: 347 ; Ceballos 1959: 223 (catalogued); Mingo & Compte 1963: 88 (♂, commented); Invrea 1964: 240 (♂); Erlandsson 1974: 31 (♂); Eiroa & Novoa 1985: 20 (♂); Pagliano & Strumia 2007: 96 (♂); Baldock 2014: 342 (♂); Lo Cascio 2015: 559 (listed); Baldock et al. 2020a: 22 (♂); Baldock et al. 2020b: 30 (♂); Parejo-Pulido et al. 2023: 16 (♂, dorsal habitus). New synonymy.

Astomyrme suberrata : Schwartz 1984: 23 (mentioned).

Paramyrme pusilla : Lelej 1984: 106 (♀, key), 113 (♀).

Physetopoda pusilla : Lelej 1985: 203 (♀; key); Lelej 2002: 61 (catalogued); Lelej & Schmid-Egger 2005: 1530 (♀); Pagliano & Strumia 2007: 79 (♀?, nec ♂?; listed); Amini et al. 2014: 590 (mentioned); Baldock 2014: 342 (♀); Lo Cascio 2015: 555 (listed); Baldock et al. 2020a: 21 (♂ [sic], ♀); Pagliano et al. 2020: 181 (catalogued); Parejo-Pulido et al. 2023: 16 (♀, dorsal habitus).

Smicromyrme (Astomyrme) suberrata : Lelej 2002: 65 (catalogued).

Astomyrme suberrata : Pagliano et al. 2018: 143 (♂, key).

Smicromyrme (Astomyrme) suberratus : Pagliano et al. 2020: 188 (catalogued); Matias 2023b: 78 (♂, key).

Diagnosis. MALE ( Figs 4A–K ). Distinguished from other Smicromyrme by combination of mandible structure (lacks external tooth basally, apically tridentate), wing venation (second abscissa of radial vein thickened, second transverse-cubital vein with spurious vein, forming “T”-shape), large vitreous and globose ocelli (diameter of anterior ocellus larger than F1 width), red propodeum, and metasoma with fringes of sparse pale pubescence. Body length 4.8–9.2 mm. FEMALE ( Figs 5A–L ). This form is distinguished from other Iberian females by the combination of the following features: mandible with basal portion short, distal element externally straight with short and proportionately broad apical tooth, subapical acute denticle and second internal acute denticle at ca. mid-length of distal element (distance between both denticles greater than apical tooth length); vertex and frons with dark recumbent pubescence only; scutellar scale very small (narrow, short), narrowed apicad, well defined; pygidium ovate with pattern of striae forming concentrical arches except over shiny apex; T2 with pale pubescence forming three basal oval spots, lateral ones small (may be insipient), and apical band medially enlarged triangularly; mesosoma all ferruginous-red with humeral angle typically not rounded (frequently subacute); antennal tubercle ferruginous-red; antenna typically dark-brown with scape apically and pedicel reddish to orange-brown; legs mostly dark, with paler orange-brown tarsi; head rounded; T2 with sub-parallel to weakly convex sides and somewhat flat dorsum. Body length 3.4–6.1 mm.

Redescription. MALE ( Figs 4A–K ). Body length 4.8–9.2 mm. Colouration. Head black, except for subapical ferruginous-red band on mandible and dark ferruginous-red antennal tubercles, dorsally with sparse brownish pubescence; mandible baso-externally with sparse yellowish-white pubescence.Antenna dark brown, ventrally paler orange-brown; scape darker, with sparse short erect whitish setae. Mesosoma ferruginous-red with black sternum, with sparse whitish erect pubescence. Wings hyaline, slightly infuscate outside the cells and inside the radial cell; veins dark-brown to black, yellowish at the wing base. Legs black with browner articulations, ferruginous-brown protibia and brown tarsi, covered with sparse white erect pubescence, denser on tibiae; yellowish-white spurs. Metasoma black, with sparse whitish pubescence forming apical fringes of recumbent pubescence on T2–3, and suberect pubescence covering T3–7 (somewhat denser on T7); S2–8 weakly fringed with whitish pubescence, more noticeably on S2–3; felt-line on T2 ca. 0.4 × T2, felt-line on S2 ca. 0.3 × felt-line of T2, composed of brown setae. Shape, structure and punctation. Head sub-elliptical, with rounded posterior angles, slightly narrower than pronotum, sparsely punctate, shiny, the punctures organized in approximately longitudinal parallel lines; clypeus with straight anterior margin, medially ecarinate (except for very short basal section), densely punctate with impunctate to weakly punctate, flat to weakly concave, shiny medial triangular section; weak subapical tubercle may be formed on each angle of the medial triangular area, with medially weakly concave subapical area; eye very large with deep internal notch; ocelli very large and convex, vitreous, anterior ocellus notably wider than diameter of F1, POL ca. 1.96 × OOL; mandible apically tridentate, with no external tooth basally. Antenna with F1 ca. 2 × pedicel and ca. 0.65 × F2 and F3. Mesosoma ovoid, pronotum dorsally short, with sub-parallel sides, rounded anterior angles and convex anterior border. Forewing with radial cell short, apically acuminate to sub-truncate; radial vein notably thickened where it meets the second tranverso-cubital vein, which is “T” shaped and thickened; pterostigma black, opaque, large (as long as costal margin of radial cell). Tegula sparsely and shallowly punctate, shiny, relatively small and convex. Tibiae unarmed. T1 short and relatively broad, apical width ca. 4.0 × medial length and ca. 0.6 × T2 maximum width, the transition to T2 with weak constriction. T2 maximum width ca. 1.45 × T2 medial length. T2 sparsely punctate over disc, punctures somewhat superficial, small and round (larger than on head), larger and less round laterally; interspaces shiny, as wide as ca. 1–3 punctures. T7 and S8 apically subtruncate, densely punctate all over, the punctures smaller on S8. Genitalia ( Figs 4I–K ). Apex of penial valves projects beyond inner protuberance of gonostylus; volsella basally slightly anterior to inner protuberance of gonostylus, apically slightly longer than apex of penial valves; volsella basally broader (lateral view), proportionately short, approximately straight, apically abruptly narrowed to a point, with row of sparse setae on dorsal margin.

FEMALE. Redescription of holotype ( Figs 6A–E ). Body length ca. 4.3 mm. Colouration. Head black, except for orange-brown antennal tubercle, ferruginous-red basal two thirds of mandible, ferruginous-brown clypeus, antennal scrobe and lower frons, and brownish-tinged gena; mandible externo-basally with few pale brownish erect setae; frons and vertex covered with moderately sparse recumbent brown pubescence and sparse erect setae of the same colour to slightly paler brown; clypeus with long pale yellowish-brown setae; lower gena with sparse whitish pubescence. Antenna dark-brown, except for (apically paler) orange-brown scape and pedicel and weakly paler reddish-brown F1–3; scape with short pale brownish to whitish sparse pubescence, with some minute setae over pedicel and F1–2; left antennae with F9–11 missing, right antenna with F11 missing. Mesosoma dorsally and laterally ferruginous-red, weakly darker brown on lower mesopleuron; pronotum antero-medially with weakly darkened interspaces; mesosomal dorsum sparsely covered with golden to orange-golden recumbent pubescence, paler over pronotal angle and pronotal spiracle; short erect dark setae sparsely over dorsum and longer on propodeum; sparse whitish recumbent pubescence on mesopleuron and pronotum laterally; propleuron brown. Legs ferruginous-brown, with femora weakly darker and tarsi paler orange-brown; procoxa dark-brown; coxae, femora and tibiae covered with long yellowish-white pubescence; meso and metatibial external spines brown; meso and metatibial spurs pale yellowish-brown. Metasoma black (S1 not observed); T2–5 and S4–5 basally browner; pygidium shiny brown; pubescence dark-brown and sparse (punctation visible), except for the following areas covered with silvery-golden appressed pubescence: T2 basomedially with ellipsoidal dense spot, basolaterally with sparse pubescence (dorsally visible as weak lateral pale spot, narrow along the tergal contour), apically with narrow fringe medially enlarged triangularly, T3 with full dense band; T6 with long pale golden to brownish-yellow (some whitish) setae laterally, partially overlapping the pygidium surface; dark-brown erect setae sparsely over areas with dark pubescence; S2 covered with moderately sparse whitish recumbent pubescence; apical fringes of sparse pale pubescence on S2–5; felt-line on T2 composed of tightly set short yellowish-brown to golden setae, overlapped anteriorly by sparse whitish pubescence part of the lateral spots. Sting partially extruded, pale brownish yellow. Shape, structure and punctation. Head. Head dorsally rounded, with short convergent sides behind the eyes and rounded posterior angles, convex posterior border, measured behind the eyes approximately as broad as pronotal anterior margin. Eye laterally prominent (dorsal view), head ca. 1.1 × broader over the eyes than behind eyes. Frons and vertex densely punctate-reticulate, punctures small round to suboval; lower frons with medial impuctate short line; narrow interspaces, some obliterate (mostly on those transverse), creating weak impression of longitudinal rows of punctures; gena densely punctate, punctures small with interspaces mostly narrower than puncture diameter, somewhat shiny. Clypeus short, dorsally micropunctate, with weakly sinuate medial transverse carina (with small tubercles along margin) and tubercle medially on upper half. Antennal scrobe punctulate, with distinct and well developed arcuate-ondulate dorsal carina from antennal tubercle to mandible base. Mandible with apex short (unidentate and weakly convex, somewhat worn), internally with one small acute subapical tooth (second denticle obscured); distal element with external margin straight; mandibles closed in this specimen. Antenna: pedicel 0.77 × F1, F1 1.08 × F2 and F1 1.08 × F3. Mesosoma. Dorsally subrectangular, length of pronotal anterior border to scutellar scale ca. 0.95 × maximum mesosomal width; pronotal anterior border weakly convex, with subparallel pronotal sides, weakly narrowed behind pronotal spiracle and weakly convex propodeal sides; humeral angle well marked, with low humeral vertical carina; propodeum with short dorsal face and subtruncate posterior face; mesosomal ratios (pronotal angle: pronotal spiracles: narrowest point: propodeum at scutellar scale level) are 93: 100: 92: 93; dorsum cracked centrally, at point of pin insertion, with right half partially lifted. Scutellar scale small and weakly protruding at a low angle (lateral view). Dorsum and upper propodeal posterior face punctate to foveate-reticulate; interspaces very narrow, some obliterate. Legs. Coxae, femora and tibiae with sparse setiferous small punctures. Metasoma. T1 short, dorsal length ca. 0.22 × apical width. T2 basally narrow, smoothly blending with T1, width ca. 2.0 × T1 apical width; densely punctate to punctate-reticulate, punctures small and somewhat elongate; interspaces mostly narrower than one puncture wide, shiny. Pygidium ( Fig. 6C ) elongate ovate, covered in subparallel striae defining concentrical arches, poorly marked medially and apically; basally partially hidden by S 5 in this specimen. Felt-line ca. 0.3 × T2 lateral length. Ratios of head width over eyes: head width behind eyes: pronotal anterior border: pronotal spiracles: mesosomal narrowest point: propodeum at scutellar scale level: T1 apical width: T2 maximum width are approximately 75: 69: 70: 76: 70: 71: 55: 100.

Type material. Holotype ♀ (photographs, MfN; Figs 6A–E ). SPAIN. CÁDIZ: Puerto Real, July 1929 (J. Waltl leg.; details in Waltl 1835), Museum für Naturkunde Berlin ( MfN, Germany), NURI: http://coll.mfn-berlin. de/u/c2e8c7. Labels ( Fig. 6E ; verbatim): Andalusien Waltl L./6438/Type [red label]/pusilla [unreadable]/ pusilla Kl det. Bischoff/ HOLOTYPE Mutilla pusilla Klug Teste B. Petersen, 1986 [red label]/ Smicromyrme pusilla (Klug) B. Petersen det. 1986/ http://coll.mfn-berlin.de/u/c2e8c7.

Material examined (117♂, 134♀). PORTUGAL (116♂, 134♀) ; all specimens R. Matias leg., det. & coll., unless otherwise stated). BRAGANÇA: Serra da Nogueira, ca. 1.5 km SW of Rebordãos ( Bragança ; alt. 974 m) : 5.vii.2018 (1♀); 7.vii.2018 (1♀). Camping park of Miranda do Douro ( Miranda do Douro ) , 15.viii.2022, 2♂ (light). VISEU: Serra do Buçaco, ca. 1.3 km SE Trezói ( Mortágua ) , 19.viii.2009, 1♂, Malaise trap (M. Matos leg., R. Matias det.), MNHNCENT0050653. LEIRIA: Cumeira ( Porto de Mós ; UTM10×10 ND08 ) , 15.x.2001, 1♂ (Bivar Sousa leg., R. Matias det.), MNHNCENT0041667. CASTELO BRANCO: Alfrívida ( Vila Velha de Ródão ) , 31.vii.2019 (2♂, one at light; 1♀). PORTALEGRE: Barragem de Montargil ( Ponte de Sôr ) , 21.iii.1973, 1♀ (P. Mendoça leg., R. Matias det.), MNHNCENT0035760. Montargil ( Ponte de Sôr ) , 9.ix.1979, 1♂ (N. Mendoça leg., R. Matias det.), MNHNCENT0035764. Santo Amaro ( Sousel ) , 16.vii.1998, 1♂ (A. Zuzarte leg., R. Matias det.), MNHNCENT0041666. SANTARÉM: Moita Redonda, Fátima ( Ourém ) : 20.vii.1974, 1♂ (N. Mendoça leg., R. Matias det.), MNHNCENT0035763 ; 22.vii.1974, 1♂ (P. Mendoça leg., D.W. Baldock det.), MNHNCENT0032380; 24.vii.1974, 1♂ (N. Mendoça leg., D.W. Baldock det.), MNHNCENT0032360; 25.vii.1974, 1♂ (P. Mendoça leg., D.W. Baldock det.), MNHNCENT0032385; 6.ix.1976, 1♂ (P. Mendoça leg., R. Matias det.), MNHNCENT0035762 ; 28.viii.1978, 1♂ (M.C. Mendoça leg., R. Matias det.), MNHNCENT0035761 ; 30.vii.1987, 1♂ (M.C. Mendoça leg., R. Matias det.), MNHNCENT0035750. Quinta das Chantas, Alcanhões , Ribeira de Santarém ( Santarém ) : 1.v.1977, 1♀ (N.C. Almeida leg., M. Romano det. 2014), MNHNCENT0032370 [specimen ex-‘ Mendoça collection’, now MNHNC, included in Baldock et al. 2020a as male P. pusilla erroneously] ; 11.vii.1977, 1♀ (P. Mendoça leg., R. Matias det.), MNHNCENT0035758 ; 29.iv.1979, 1♀ (P. Mendoça leg., R. Matias det.), MNHNCENT0035759 ; 19.viii.2001, 1♂ (P. Mendoça leg., D.W. Baldock det.), MNHNCENT0032382. Santarém ( Santarém ) , 25.vi.1968, 1♂ (P. Mendoça leg., R. Matias det.), MNHNCENT0032485. Vasa-Sacos saltpans, RNET ( Benavente ) , 16.xi.2017, 1♀. LISBON: Casal Palear—Casal do Sol (Carvoeira, Torres Vedras ) : 13.viii.2019 (1♀); 7.ix.2019 (1♀); 10.ix.2019 (1♀); 3.v.2021 (1♀); 26.v.2021 (1♀); 7.vii.2021 (1♀); 26.viii.2023 (1♀); 28.viii.2023 (1♀); 6.ix.2023 (1♀); 14.ix.2023 (1♀). Casal Sobrigal (Carmões, Torres Vedras ) : 24.viii.2001 (1♂, light); 5.ix.2001 (1♂, light); 4.vii.2004 (1♀); 31.vii.2005 (1♂, light); 4.ix.2005 (2♂, light); 7.ix.2005 (1♂, light); 3.vii.2021 (1♂, light); 23.vii.2021 (1♀; pale green pan trap); 28.vii.2021 (3♂); 29.vii.2021 (1♂, light); 30.vii.2021 (1♂, light); 2.viii.2021 (1♂, 16:40GMT; 4♀); 4.viii.2021 (5♂: one in white pan trap, one in blue pan trap, two in light green pan trap, one at light); 5.viii.2021 (4♀); 6.viii.2021 (3♀); 8.viii.2021 (2♂: one in white pan trap, one in yellow pan trap; 1♀); 9.viii.2021 (3♀); 10.viii.2021 (1♀); 11.viii.2021 (2♂, light); 12.viii.2021 (3♂, light; 1♀); 30.ix.2021 (1♀); 7.v.2022 (1♀); 21.vi.2023 (1♂; light); 23.vi.2023 (2♂; light); 28.iv.2023 (1♀); 30.iv.2023 (1♀); 2.v.2023 (1♀); 4.v.2023 (2♀); 5.v.2023 (1♀); 23.vi.2023 (1♀); 24.vi.2023 (2♀); 17.viii.2023 (2♂; light); 18.viii.2023 (2♂; light); 20.viii.2023 (2♂; light); 21.viii.2023 (1♂; light); 22.viii.2023 (1♂; light); 15.ix.2023 (1♂; light). Carrasqueira (Carmões, Torres Vedras ) , 26.iv.2023, 2♀. Carmões (Carmões, Torres Vedras ) : 2.v.2017 (2♀); 11.iii.2020 (4♀); 4.x.2021 (1♀); 7.x.2021 (3♀). Quinta do Hespanhol (Dois Portos, Torres Vedras ) : 29.vii.2000 (3♀); 5.v.2002 (2♀); 25.viii.2002 (3♀); 11.ix.2002 (4♀); 6.x.2002 (1♀); 1.vi.2003 (2♀); 29.vii.2003 (2♀); 2.v.2019 (2♀); 20.v.2021 (4♀); 21.v.2021 (2♀); 31.v.2021 (1♀). Quinta Nova do Hespanhol (Carvoeira, Torres Vedras ) : 4.ix.2001 (1♀); 10.vi.2002 (2♀); 7.vii.2002 (1♀); 3.viii.2002 (2♀); 31.vii.2003 (9♀); 2.viii.2003 (2♀); 8.iv.2017 (1♀). Quinta do Bouro (Pereiro de Palhacana, Alenquer ) , 5.v.2023, 1♀. Ericeira (Ericeira, Mafra ) , 22.v.2019, 2♀. Foz do Lizandro (Ericeira, Mafra ) , 28.vi.2019, 5♀. Mafra-Gare ( Mafra ) , 31.viii.2020, 1♀. Azenhas do Mar ( Sintra ) : 11.vii.2019 (1♀); 22.vii.2020 (1♀). Serra do Lima (Odrinhas, Sintra ) , 31.iii.2017 (1♀); 24.iv.2018 (1♀); 22.vi.2018 (1♀); 16.iv.2019 (3♀). Serra das Éguas (Carenque, Amadora ) : 11.vi.2019 (2♀); 15.vii.2020 (1♀); 10.viii.2020 (1♀); 15.vii.2022 (1♀). FCUL, Campo Grande ( Lisbon ) , 14.x.2002, 1♀ (I. Silva leg., R. Matias det. & coll). Cascais ( Cascais ) , 28.vi.1983, 1♂ (P. Mendoça leg., R. Matias det.), MNHNCENT0032392. SETÚBAL: Serra da Arrábida ( Setúbal ) , 19.vii.1978, 1♂ (A. Serrano leg., R. Matias det.), MNHNCENT0000863. Galo quarry (Zambujal de Cima, Sesimbra ) , 30.v.2002, 1♀. Cape Espichel (Azóia, Sesimbra ) , 25.iv.2008, 1♀. Facho de Santana ( Sesimbra ) , 20.iv.2019, 1♀. Parque Luso (Corroios, Seixal ) , 23.v.2022, 2♀. ÉVORA: Herdade da Lobeira ( Montemor-o-Novo ) , 30.vii.1997, 2♂ (A. Zuzarte leg., R. Matias det.), MNHNCENT0041665. Vale das Quintas, Veiros ( Estremoz ) , 9.vii.1998, 1♂ (A. Zuzarte leg., R. Matias det.), MNHNCENT0041664. Alpista ( Montemor-o-Novo ) , 1.iv.2001, 1♀ (J.C. Rodrigues leg., R. Matias det. & coll). BEJA: Vila Nova de Milfontes ( Odemira ) , 11.vii.1972, 1♂ (C. Mendoça leg., D.W. Baldock det.), MNHNCENT32271. Herdade da Coitadinha, Parque de Natureza de Noudar ( Barrancos ) , 5.vi.2023, 1♂ ( T. Moreira leg., R. Matias det.), MNHNCENT0050652. FARO: Sagres ( Vila do Bispo ) , 15.ix.2000, 1♀ (I. Silva leg., R. Matias det. & coll). Sesmarias ( Albufeira ) , 20.v.1995, 1♂ (D. Keen leg., M. Romano det.), MNHNCENT0041628. RIAS (wildlife rehabilitation and research center), ca 1 km East of Olhão (Quelfes, Olhão ) : 9.vii.2022 (7♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.) ; 10.vii.2022 (8♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.) ; 22.vii.2022 (5♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.) ; 28.vii.2022 (31♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.). Silves , 12.x.2010, 1♂ (D.W. Baldock leg., M. Romano det.), MNHNCENT0041627. Arrochela (ca. 200 m S of Arade River), Silves : 15.vii.2022 (1♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.) ; 16.vii.2022 (1♂), light trap ( T. Valkenburg leg., R. Matias det. & coll.) . SPAIN (1♂) . ALMERIA: Los Moras ( Adra ) , 12.vii.2023, 1♂, light trap (F. Rodriguez leg, R. Matias det. & coll.) .

Intraspecific variation ( Figs 5A–L , 7–10 , 19A–D , 21A–D ). FEMALE (N = 114). Body length 3.4–6.1 mm. Colouration. Head ( Figs 7A–F ): integument most frequently black, some specimens browner as holotype; sparse recumbent pubescence on frons and vertex (n = 107) most frequently black (n = 81), some specimens (n = 9; e.g. Figs 7B, 7E ) with browner pubescence as holotype, dark-brown (n = 9), reddish-brown (n = 6) or yellowish-brown (n = 2); erect setae black (most frequently) to pale-brown; occasionally, reddish-brown pubescence replaces all black pubescence on head and metasoma; some specimens (n = 16) with pubescence of head and elsewhere extremely abraded, with few fragments of setae remaining. Clypeus ferruginous-red to brown, with colour variably extending to antennal scrobe, malar space and lower frons. Antenna (N = 111) somewhat variable in colour (within each population): most frequently (n = 100) brown to dark-brown, except for darker (dark-brown to blackish-brown) scape (apically paler reddish), orange to reddish-brown pedicel and F1–3 (as holotype); less frequently (n = 10) antenna pale ferruginous-orange to orange-brown, paler on pedicel and ventrally on F1–8, and rarely (n = 1) antenna all dark-brown, with blackish-brown pedicel and F1. Antennal tubercle ferruginous-red. Mesosomal dorsum ( Figs 8A–I ) covered with variable proportion of recumbent golden setae and finer black setae: some specimens with golden setae only (similar to holotype), most with a proportion of black setae; golden setae frequently reduced to lateral spot over pronotal spiracle (e.g. Figs 5K , 8H–I ). Pronotum most frequently ferruginous-red without darkening; some specimens with variably weakly darkened interspaces medially and one specimen with narrow darkened band across anterior pronotal margin; sparse sub-recumbent black setae frequently over darkened areas. Scutellar scale (N = 111) concolourous with metasomal dorsum to weakly darkened (n = 30), or apically blackish (n = 81). Legs variable as antenna, with insects with darkest antenna most frequently also darker on legs; in a sample of 111 specimens, most (n = 63) medium reddish-brown, weakly darker brown on femora and tibiae apically, and paler orange-brown tarsi, procoxa blackish-brown (similar to holotype); darker specimens also frequent (n = 34), with darker brown areas replaced by blackish-brown (mainly femora); paler specimens less frequent (n = 13), where legs may be uniformly orange-brown, sometimes also on procoxa; one specimen exceptionally pale, with legs uniformly yellowish. Metasoma ( Figs 9A–I ) mostly black, sometimes with browner integument, except for T1 , S1 and S6; T1 (N = 99) most frequently dark-brown (n = 62), also blackish (n = 19) or dark-red (n = 16), rarely ferruginous-red (n = 2); S1 (N = 90) most frequently dark-red with blackish longitudinal medial carina (n = 50), to dark-brown (n = 26), less frequently blackish (n = 28); S6 (N = 95) most frequently brown (n = 63) or orange-brown (n = 29), rarely reddish (n = 2) or black (n = 1). T2 with baso-lateral spots of pale pubescence variable in density, from well defined spots (e.g. Figs 9A–C ) to obscure in dorsal view (e.g. Fig. 9I ); medial spot variable in shape and relative size; apical band slightly variable in width and medial shape, sometimes somewhat abraded. Pygidium (N = 107; Figs 10A–L ) most frequently dark-brown (n = 49) to black (n = 31), less frequently paler brown (n = 28); laterally always margined by sparse pale setae, normally all golden, sometimes yellowish-white; setae may be partially abraded. Shape, structure and punctation. Head shape and punctation approximately as in holotype, interspaces most frequently shiny, occasionally duller. Mandible (N = 109) somewhat worn on most specimens (n = 49), extremely worn (less than basal ca. 60% remaining) on 16 specimens, and with little to no abrasion visible on 44 specimens, where the complete structure is observable: apical tooth acute, short and broad, externally convex, followed by small subapical acute denticle, with second similar denticle (visible in dorsal view) at approximately half length of the inner margin of the distal element (which is narrow and externally straight), both denticles separated by gap wider than apical tooth length (see Discussion for illustrated comparison of this feature with other Iberian Smicromyrmini : Fig. 21 ). Clypeus with similar structure across the series (and holotype; frontal view), short, subtruncate (dorsal view); notable longitudinally elongate tubercle medially on upper half (visible on dorsal view); lower half weakly concave variably angled posterad; anterior margin of medial transversal carina variably tuberculate, frequently with one small tubercle on each side anterolaterally, defining a low triangle together with the medial tubercle of the upper half. Hypostomal carina well developed. Mesosoma ( Figs 8A–I ) varies slightly in proportions, keeping the same general structure, although some specimens have particularly well marked humeral angles and are flatter on dorsum; most specimens have sub-rectangular mesosoma, some sub-quadrangular; mesosomal length to scutellar scale 0.94– 1.09 × maximum mesosomal width; sides of mesosoma (dorsal view) weakly concave to straight. Scutellar scale (N = 109) very small though well defined, basally as narrow as ca. 1–1.5 adjacent punctures, as long or slightly shorter than it is wide, shaped as dorsally arcuate triangular lamella (more or less rounded apically, sometimes weakly acute) or short blunt tubercle; rarely (n = 2) scale slightly wider (ca. two punctures wide basally), connected to adjacent raised interspaces with somewhat undefined shape ( Figs 5H, 5K–L , 8A–I ). Legs. Meso and metatibia externally with ca. 8–9 short spines in two rows, and 3–4 finer apical spines. Metasomal shape ( Figs 9A–I ): T2 somewhat elongated, more or less convex laterally; T2 width ca. 0.96–1.07 × T2 length; on some specimens T2 proportionately broader and shorter; more or less flattened on disk, typically laterally strongly curving downwards and slightly inwards. S6 apically bidentate, apicomedially concave. T2 densely punctate across the series, punctures small most frequently slightly elongate; one specimen with weakly impressed punctures on disc, almost impuctate. Pygidium ( Figs 10A–L ) elongated, shape varies from sub-oval to narrow ovate, sometimes approaching a deltoid shape, length 1.3–1.9 × maximum width; apex and sometimes medial area shiny, devoid of striae, but in some specimens these areas are also striated (e.g. Figs 10D, 10G, 10H ); laterally weakly carinate on the apical third. S1 with longitudinal medial carina, rarely forming low tooth anteriorly .

MALE (N = 80). Body length 4.8–9.2 mm. Colouration. Propodeum wholy red on most specimens (n = 60), with a smaller proportion weakly darkened basally on the posterior face (n = 14), or browner on the posterior propodeal face and over the metapleuron (n = 6), never black. Mesopleuron most frequently either black over its lower half (n = 32) or slightly less extensively (n = 31); less frequently black reduced mostly to ventral surface (n = 8) or vestigial on ventral surface (n = 6), rarely all red (n = 3); on some specimens (n = 7) dark integument browner. Antenna variable, frequently extensively yellowish-brown on the ventral surface, rarely fully brown. Shape, structure and punctation. POL> OOL. T1 short and variably broad, apical width ca. 2.7–5.3 × medial length (dorsal view) and ca. 0.6–0.7 × T2 maximum width, smoothly blending with T2 or with weak constriction. T2 maximum width ca. 1.34–1.60 × T2 medial length. Punctation of T2: punctation always denser and larger laterally than over disc, most frequently with interspaces of one puncture wide or less; density and diameter of punctures over disc variable, with punctures most frequently rather small, and average interspaces varying from ca. one to about four punctures (from disc) wide; most specimens (n = 67) had average interspaces of 1.5–2 times the diameter of a puncture, less frequently with interspaces narrower (n = 5; ca. one puncture wide), or wider (n = 8; ca. 2.5–4 times the diameter of a puncture); the interspaces are most frequently also variable on each specimen, i.e. punctures somewhat irregularly distributed. Two specimens show bidentate mandible, apparently due to abrasion of the middle tooth. One specimen has notably proportionately smaller, though otherwise normal, head. The specimen from Almería ( Spain) fits within the variation above described, both in terms of colour and structure, presenting average features (see slso the section for S. opistomelas , below, for further discussion of some of these features).

Original description [Klug in Waltl 1835: 92]. Mutilla usilla [sic] Klg. Nigra, thorace rufo, abdomine maculis 3, lateralibus obsoletis fasciaque postica argenteis. [Following text translated from German, some terms adapted.] A single female, about the size of a small M. rufipes . The body is fairly densely punctate all over, the basic colours black. On the head there is no trace of white patches or similar pubescence. The area under the antennae, antennal tubercle, and the base of the mandibles are red. The antennae are blackish except for the tip of the first segment [scape], which is red. The mesosoma is almost quadrangular, monochromatic brown-red, with almost imperceptible silver-grey pubescence dorsally, long blackish setae laterally. The legs are blackish, gray-haired, with reddish tarsi; metasoma grey-haired. At the base of the second metasomal tergum there is a round, glossy, silvery-white spot in the middle; towards the side, more downwards in the same direction, there is also a similar spot. The second and third metasomal segments are broadly covered in silver-white pubescence on the apical margin, forming a transverse band. The metasomal sternal segments, especially the medial ones, are fringed with whitish hair.

Distribution. According to literature, reported from the following areas. Europe: Portugal ( Figs 11A–B ), Spain, France (south, and Corsica: females only), Italy (Piedmont, Liguria, Emilia-Romagna, Tuscany, Lazio, Apulia, Basilicata, Calabria; also Sicily, Sardinia, Lampedusa), Malta; also reported from Slovakia, Hungary, Romania, Albania, Croatia, Bulgaria, Greece. Asia: Cyprus, Turkmenistan? North Africa: Morocco, Algeria, Tunisia, Libya, Egypt (e.g. Lelej 2002; Bogusch 2006; Pagliano & Strumia 2007; Muskovits & György 2011; Pagliano & Matteini Palmerini 2014; Pagliano et al. 2020). Males (i.e. S. suberrata Invrea ) only recorded from Portugal, Spain, Southern France, southern Italy (Calabria), Lampedusa Island ( Italy), Malta, Morocco, Algeria, Libya and Egypt). This extensive distribution may in fact congregate more than one species; see Discussion ( Fig. 20 ).

Remarks. A) Holotype. Some of the reddish and darker tones of the holotype, mentioned in the original description (Klug in Waltl 1835; see above) are apparently paler (orange to brownish) in the photographs of the specimen, perhaps due to ageing.

B) Behavioural notes. Males are generally found at night, atracted to artificial lights, but most other behaviour of the species is unknown. At Casal Sobrigal males were occasionally found during the day: one was collected at 16:40 GMT, 2.viii.2021, walking with wings raised on dry mossy and grassy ground under a stand of false yellowhed ( Dittrichia viscosa ), where females had been collected on other occasions; another male was observed for some seconds, before 19:00 GMT (8.viii.2021), though not collected, perched on dry grass ca. 20 cm above soil, close to the previous point. Specimens were collected inside pan traps at early dusk on 4.viii.2021: one inside a blue pan trap, two inside a light green trap and one inside a white trap between 19:50 and 20:00 GMT; one of the previous was found already inside the trap, whilst the other three were observed flying in the immediacy of the pan traps before falling inside, possibly prospecting for females (close to the first mentioned point, above). Finally, two males were retreived from pan traps at 14:00 GMT that must have fallen during the late morning hours (one of the males, found motionless, fully recovered less than one hour after drying). These anecdotal observations suggest the male of the species is not exclusively nocturnal, despite being most easily found at night.

C) Additional data. Diagnosable photographic records of the species (males only, whenever the identity of the species could be safely established) divulged in the citizen science website iNaturalist (www.inaturalist.org; last accessed 8.vi.2024) were also considered for defining the distribution of the species in Portugal ( Fig. 11A ). [Diagnosing features: wing venation as in Fig. 4H , mesosoma all red, metasoma with pale pubescence restricted to sparse apical fringes, ocelli large; mandible structure most frequently not observable. Observations of which exact locality has been obscured by the original recorder have not been considered.]. PORTUGAL. VILA REAL. ca. 250 m SE Brunhedo (Ribeira de Pena; 41°30'57.6"N 7°47'03.3"W; 475 m asl; UTM10×10 PF09 ), 26.vii.2022, 1♂ (P. Alves photo; https://www.inaturalist.org/observations/128058816) . LEIRIA. Casal Novo (Amor; 39°46'46.6"N 8°52'29.5"W; 56 m asl; UTM10×10 NE10 ), 28.vii.2023, 1♂ (S. V. Duarte photo; https://www.inaturalist.org/ observations/175580370) . GUARDA. São Paio ( Gouveia ; 40°30'51.4"N 7°35'17.7"W; 550 m asl; UTM10×10 PE28 ), 8.viii.2023, 1♂ (‘brandao’ photo; https://www.inaturalist.org/observations/186058478) . CASTELO BRANCO. Espinho Grande (Proença-a-Nova; 39°43'43.6"N 7°50'40.6"W; 308 m asl; UTM10×10 ND99 ), 29.vii.2022, 1♂ (R. Martins photo; https://www.inaturalist.org/observations/128678710) . PORTALEGRE. Vale de Carvão (Santo António das Areias; 39°23'39.7"N 7°20'08.9"W; 464 m asl; UTM10×10 PD46 ), 9.vii.2023, 1♂ (N. Fletcher photo; https://www.inaturalist.org/observations/171929384) . FARO. Corte do Gago (Azinhal, Castro Marim ; 37°17'23.3"N 7°32'45.7"W; 160 m asl; UTM10×10 PB22 ), 22.x.2010, 1♂ (V. Jacinto photo; https://www.inaturalist. org/observations/171453227) .

Habitat. Ubiquitous. Females have been found from sea level to ca. 1000 m asl, from urban settings to forested areas; males have been recorded also from sea level to ca. 700 m asl, with most records biased to urban habitats (surrounded by various habitats, including coastal areas, agricultural land, woodland) due to the presence of attractant artificial light, but also in open woodland (including Eucalyptus plantations) away from urbanization.

Figures

FIGURE 4. Smicromyrme pusillus (Klug in Waltl), male (=S. suberrata Invrea), Casal Sobrigal, 28.vii.2021 (RMC1375). A dorsal habitus B ventral habitus C head, frontal view D clypeus E mandible, dorsal view F mandible, lateral view G lateral habitus H wings, dorsal view I genitalia, dorsal view J genitalia, ventral view K genitalia, lateral view. Scale bar: 1 mm (A–B, G–H), 0.2 mm (I–K), 0.25 mm (D–F), 0.5 mm (C).

FIGURE 5. Smicromyrme pusillus (Klug in Waltl), female, Casal Sobrigal, 5.viii.2021 (RMC1421), same specimen as in pairing trial 32 (copula). A dorsal habitus B ventral habitus C lateral habitus D pygidium E antenna F mandible, ventral view G head, frontal view H mesosoma, dorso-lateral view I clypeus J metasoma, felt-line, lateral view K mesosoma, dorsal view L scutellar scale (white arrow). Scale bar: 0.5 mm (A–C), 0.1 mm (D, I, L), 0.4 mm (E), 0.2 mm (F), 0.25 mm (G–H, J–K).

FIGURE 6. Smicromyrme pusillus (Klug in Waltl), holotype, female (July 1829, Puerto Real, MfN; NURI: http://coll. mfn-berlin.de/u/c2e8c7). A dorsal habitus B head, frontal view C pygidium D lateral habitus E labels. Scale bar: 0.5 mm (A), 0.25 mm (B), 0.1 mm (C), 1 mm (D). Photographs reproduced with permission of MfN.

FIGURE 7. Smicromyrme pusillus (Klug in Waltl), female, head, dorsal view, variation. A Carmões, 11.iii.2020 (RMC1099) B Serra do Lima, 16.iv.2019 (RMC639) C Serra do Lima, 16.iv.2019 (RMC643) D Carmões, 11.iii.2020 (RMC1100) E Foz do Lizandro, 28.vi.2019 (RMC783) F Casal Sobrigal, 2.viii.2021 (RMC1409). Scale bar: 0.25 mm (A–F). Note 5E shows dorsal view aligned with occiput, affecting exact shape of head.

FIGURE 8. Smicromyrme pusillus (Klug in Waltl), female, mesosoma, dorsal view, variation. Note different proportions and lateral shape (weakly concave to straight); pale pubescence on dorsum extensive on A–C (B unusually silvery), reduced to lateral areas and over mesoscutum on D–F, and further reduced to vestigial spots over the pronotal spiracles on G–I; pronotum may be weakly and diffusely darkened medially on interspaces. A Carmões, 11.iii.2020 (RMC1099) B Serra do Lima, 16.iv.2019 (RMC639) C Quinta do Hespanhol, 7.vii.2002 (RMC260) D Carmões, 11.iii.2020 (RMC1100) E Foz do Lizandro, 28.vi.2019 (RMC783) F Casal Sobrigal, 2.viii.2021 (RMC1409) G Casal Sobrigal, 5.viii.2021 (RMC1422) H Quinta Nova do Hespanhol, 10.vi.2002 (RMC251) I Serra do Lima, 16.iv.2019 (RMC643). Scale bar: 0.25 mm (A–I).

FIGURE 9. Smicromyrme pusillus (Klug in Waltl), female, metasoma, dorsal view, variation. Note variability of shape of T2 (basally and laterally), and shape, density and extension of the three pale pubescent basal spots and apical fringe of T2. A Carmões, 11.iii.2020 (RMC1099) B Serra do Lima, 16.iv.2019 (RMC639) C Quinta do Hespanhol, 7.vii.2002 (RMC260) D Carmões, 11.iii.2020 (RMC1100) E Foz do Lizandro, 28.vi.2019 (RMC783) F Casal Sobrigal, 2.viii.2021 (RMC1409) G Casal Sobrigal, 5.viii.2021 (RMC1422) H Quinta Nova do Hespanhol, 10.vi.2002 (RMC251) I Serra do Lima, 16.iv.2019 (RMC643). Scale bar: 0.25 mm (A–I).

FIGURE 10. Smicromyrme pusillus (Klug in Waltl), female, pygidium, variation. A Carmões, 11.iii.2020 (RMC1099) B Serra do Lima, 16.iv.2019 (RMC639) C Quinta do Hespanhol, 7.vii.2002 (RMC260) D Casal Sobrigal, 5.viii.2021 (RMC1422) E Foz do Lizandro, 28.vi.2019 (RMC783) F Carmões, 11.iii.2020 (RMC1100) G Casal Sobrigal, 2.viii.2021 (RMC1409) H Quinta Nova do Hespanhol, 10.vi.2002 (RMC251) I Serra das Éguas, 15.vii.2022 (RMC1676) J Quinta Nova do Hespanhol, 31.vii.2003 (RMC352) K Quinta do Hespanhol, 11.ix.2002 (RMC318) L Carvoeira, 26.v.2021 (RMC1286). Scale bar: 0.1 mm (A–L).

FIGURE 11. Occurrence of Smicromyrme pusillus (Klug in Waltl) and S. matritensis (Mercet) in Portugal. A Smicromyrme pusillus, males (= S. suberrata Invrea; yellow dots: photographic records; see Remarks: Additional data) B Smicromyrme pusillus, females (= Physetopoda pusilla) C Smicromyrme matritensis (red dot: male). Localities are shown on UTM 10×10 km grid.

FIGURE 19. Comparison between male Smicromyrme pusillus (Klug in Waltl) (A–D) and S. opistomelas Invrea (E–F); mesosoma, dorsal view (left), T1–2, dorsal view (right).A–B Smicromyrme pusillus, Casal Sobrigal (Torres Vedras, Portugal), 28.vii.2021 (RMC1375) C–D Smicromyrme pusillus, Almería (Spain), 12.vii.2023 (RMC2093) E–F Smicromyrme opistomelas, Almería, 12.vii.2023 (RMC2092). Scale bar: 2.5 mm (A–F).

FIGURE 20. Distribution of Smicromyrme (Astomyrme) pusillus (Klug in Walt) and S. (A.) opistomelas (Invrea). Legend. Dots: exact localities; triangles: centre of countries or regions when exact locality unknown; stars: type localities. From left to right: P Puerto Real (Spain; type locality of Mutilla pusilla); G Gheddadia (Libya; type locality of S. suberrata); O Bu Fachra (Cyrenaica, Libya; type locality of S. opistomelas) C: Mount Troodos (Cyprus; locality for male S. pusilla sensu Bischoff 1933) A: Ashkhabad (Turkmenistan; type locality for male Mutilla subcomata sensu Radoszkowski 1888). White dots/triangles: S. pusillus, females (published data); yellow area/dots/arrows: S. suberrata sensu Invrea; blue areas/dots: S. opistomelas; red dots: putatite male Smicromyrme pusilla sensu Bischoff 1933; small yellow arrows (left to right) indicate Lampedusa Island, Malta and Pizzo Calabro (Calabria, Italy). Based on André 1910; Baldock et al. 2020a, b; Bischoff 1933; Bogusch 2006; Eiroa & Novoa 1985; Invrea 1953b, 1957, 1964; Giner Marí & Suárez 1959; Lelej 2002; Lelej et al. 2003; Muskovitz & György 2011; Pagliano & Matteini Palmerini 2014; Pagliano et al. 2020; personal data. Data not considered: Berland 1925; S. pusilla (Morocco) in Giner Marí & Suárez 1959 (misidentifications, according to Suárez); published records from Spain (due to uncertainty of specimen identity at genus level; see Matias 2024; this paper).

FIGURE 21. Mandible shape in female Iberian Smicromyrmini. A–D Smicromyrme (Astomyrme) pusillus (Klug in Waltl) [A Casal Sobrigal (Torres Vedras), 5.viii.2021 (RMC1421), lateral view: note second denticle not visible due to angle (compare with Figs 5G, 5I) B Cádiz (Spain), holotype: second denticle obscured; note abraded apical tooth C Mafra-Gare, (Mafra), 31.viii.2020 (RMC1212), dorsal view D Casal Palear—Carvoeira (Torres Vedras), 26.v.2021 (RMC1286), dorsal view] E Smicromyrme (Astomyrme) matritensis, Galicia (Spain), 14.v.2019 (RMC678), dorsal view F Ephutomma montarcense, Casal Sobrigal, 6.viii.2021 (RMC1430), dorsal view G Physetopoda cf. sericeiceps, 29.viii.2019, Fonte da Telha (RMC1039), dorsal view H Physetopoda cf. lucasii, Alfrívida (Castelo Branco), 31.vii.2019 (RMC917), dorsal view I Smicromyrme partitus, Casal Palear—Carvoeira, 25.iv.2022 (RMC1565), dorsal view J Smicromyrme ruficollis, Vila Nova de Milfontes (Beja), 17.vii.2019 (RMC848), dorsal view K Smicromyrme sulcisius, Fonte da Telha (Setúbal), 27.vi.2019 (RMC763), dorsal view. Legend. 1: first denticle; 2: second denticle. Scale bar: 0.1 mm (A–K).