Smicromyrme (Astomyrme) opistomelas Invrea, 1950

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 : 43-50

Smicromyrme (Astomyrme) opistomelas Invrea, 1950

( Figs 18A–K )

Smicromyrme subcomata var. opistomelas Invrea, 1950: 26 , ♂ (original combination), type locality: Bu Fachrà ( Cyrenaica , Libya), vi [june], G.C. Krüger leg., holotype in Museo Civico di Storia Naturale “ G. Doria ” (Genova, Italy). Invrea 1953b: 289 (♂) .

Smicromyrme subcomata : Suárez 1952: 83 (♂ in part?, nec ♀).

Smicromyrme suberrata forma opistomelas : Invrea 1957: 199 (♂, mentioned); Invrea 1964: 243 (♂).

Smicromyrme (Astomyrme) opistomelas : Lelej 1985: 212 (key); Lelej 2002: 65 (catalogued); Pagliano et al. 2020: 188 (catalogued); Matias 2023b: 78 (♂, key).

Smicromyrme opistomelas : Pagliano & Strumia 2007: 93 (♂).

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

Diagnosis. MALE ( Fig. 18 ). Recognized by combination of apically tridentate mandible lacking external tooth basally, wing with second abscissa of radial vein thickened and second transverse-cubital vein with spurious vein (forming “T”-shape), large vitreous and globose ocelli (diameter of anterior ocellus larger than F1 width), black propodeum, and metasoma with sparse pale pubescence forming apical fringes. Body length 6.5–8.0 mm. FEMALE. Unknown.

Material examined (3♂). SPAIN. (3♂; all specimens F. Rodriguez leg., R. Matias det. & coll.). ALMERIA: La Envía, Aguadulce ( Roquetas del Mar ), 9.vii.2023, 2♂ (light). Los Moras (Adra), 12.vii.2023, 1♂ (light) .

Intraspecific variation. MALE (N = 3). Body length 6.5–8.0 mm. Colouration. Propodeum fully black on two specimens, weakly browner next to the metapleural suture in one specimen. Mesopleuron black over less than its lower half (n = 2) or wholly red (n = 1; Figs 18B, 18G ); metapleuron wholly black (n = 1), browner posterad (n = 1) or red anteriorly and black on posterior half (n = 1). Punctation of T 2. Size of punctures somewhat variable in this sample, from approximately as small as in S. pusillus (= S. suberrata Invrea ) to larger; punctation denser laterally than on disk, but overall density variable: interspaces on disk vary from ca. 1–2 times puncture diameter, being denser on the specimen with larger punctures. Genitalia ( Figs 18I–K ). Not visibly distinct from those of S. pusillus .

Distribution. Europe: Spain (southeast), Italy (Sardinia). North Africa: Morocco, Libya (see map in Discussion: Fig. 20 ). Although Invrea (1964) indicates Iberian Peninsula (rather than Spain, suggesting a wider distribution), this taxon is presently not known from Portugal.

Remarks. Diagnosing criteria other than those in the short original description ( Invrea 1950) have been indicated by Pagliano & Strumia (2007), and repeated by Pagliano et al. (2018), namely the density of punctation over T2; according to those authors, S. suberrata (i.e. the male S. pusillus ) would have larger punctures, regularly distributed, with interspaces ca. 1–1.5 times the diameter of one puncture, while S. opistomelas would have smaller punctures, with wider interspaces over disc than laterally, ca. 2–3 times their diameter. Further, Pagliano & Strumia (2019) additionaly define S. opistomelas as having red or brown propodeum and linear second transverse-cubital vein, which is in strong contrast and disagreement with the original description of this taxon, in which Invrea (1950: 26– 27) is clear when stating var. opistomelas differed from a male from Mallorca island, Spain (considered to represent S. subcomata sensu Radoszkowski = S. suberrata = male S. pusillus ) only in having black propodeum and sternum; Pagliano & Strumia (2019) also add S. opistomelas would have slender general structure, which contradicts features previously indicated by the same authors to separate it from S. corriasi ( Pagliano & Strumia 2014: 62) .

The sample of Portuguese and Spanish male S. pusillus (= S. suberrata ) and S. opistomelas examined for this study does not fully confirm the above indicated criteria based on punctation to diagnose both species (e.g. Fig. 19 ). In fact, the Portuguese male S. pusillus have variable punctation density, with an overlap with the density attributed above to S. opistomelas (i.e. punctures small, with variable and wider interspaces over disc than laterally where punctures larger); on the other hand, though a small sample, the Spanish S. opistomelas had denser and larger punctation (notably in one specimen; Fig. 19E ) than almost all S. pusillus examined, fully overlapping the puncture density indicated for S. pusillus by Pagliano & Strumia (2007). These observations suggest puncture density and puncture size are unreliable as diagnostic features to separate these species, at least over a significant part of their distribution area.

A clarification, with further research, would be useful regarding the validity of S. opistomelas as an independent species, rather than potentially being part of the chromatic variation of S. pusillus ; note, however, that both forms are sympatric in Almería, and probably elsewhere (which seems to argue against the possibility that the black propodeum would simply be a phenotypic consequence of abiotic variables, such as temperature and humidity).

Synonymic notes. This taxon, elevated to species-rank by Lelej (1985), was originally described as a ‘variety’ of the male Smicromyrme subcomata (sensu Radoszkowski) by Invrea (1950), the latter renamed S. suberrata Invrea, 1957 ; opistomelas was then treated as a ‘form’ of S. suberrata ( Invrea 1957, 1964).

Should this species be considered conspecific with S. suberrata Invrea , then it would become its senior synonym, but would subsequently fall in synonymy with S. pusillus , following the results herein presented. By considering Smicromyrme subcomata var. opistomelas Invrea, 1950 a form of Smicromyrme suberrata Invrea 1957 , Invrea (1957: 199) effectively considered both taxa were conspecific, and therefore the species-group name opistomelas should have had precedence over the new name suberrata ; thus, if conspecific but subspecifically different, suberrata would be a form, or subspecies, of opistomelas , and not the other way round; Art. 23 of the Code (ICZN 1999).

The situation of the species is further confused by the recent description of a similar species in Sardinia ( S. corriasi Pagliano in Pagliano & Strumia, 2014) whose diagnosis seems to fall within the described variation of S. opistomelas and is based on subtle criteria that are known to vary slightly, sometimes markedly, within a given species (e.g. colour of integument, density of punctation, robustness); the differences indicated by Invrea (1957, 1964) separating specimens from North Africa from those of Europe (antennae paler overall in Africa, paler on the ventral side in Europe, but also on punctation density) may account for part of this variation. The available evidence seems to suggest S. corriasi may represent a junior synonym of S. opistomelas .

It is beyond the scope of this paper to determine the conspecificity of S. opistomelas and S. suberrata (= S. pusillus ), as well as of S. corriasi . In the absence of adequate data (including molecular information), S. opistomelas is thus here considered as a separate species, following the current view (e.g. Lelej 2002; Pagliano et al. 2020).

Key to Iberian Smicromyrme

FEMALES (unknown for S. ferdinandi Invrea, 1953 , S. merceti ( André, 1903) , S. metanotalis metanotalis (André, 1902) , S. metanotalis andradei Suárez, 1954 and S. opistomelas )

1. Mandible with two internal denticles (preapical denticle and second internal denticle at ca. half length of distal element) separated by distance notably longer than apical tooth length ( Fig. 21C ). Scutellar scale short, narrow and inconspicuous, but well defined (e.g. Fig. 5L ). Pygidium weakly broader apically than basally ( Fig. 10 ). Mesosoma approximately rectangular (dorsal view), with pronotal angles developed ( Fig. 8 ). [Mandible with distal element straight]....................... 2

- Mandible with two subapical internal denticles separated by distance shorter than apical tooth length (e.g. Fig. 21G ). Scutellar scale most frequently conspicuous and well developed, transverse (small in S. plantourianus ). Pygidium parallel sided or triangular and broad based. Mesosoma trapezoidal to weakly rectangular (dorsal view), with pronotal angles normally somewhat rounded. [Mandible with distal element straight or somewhat convex]........................................... 3

2. Vertex with dark pubescence only ( Fig. 7 ). T2 with lateral spots sparse, poorly defined ( Figs 5J , 9 ). Pronotal anterior margin not darkened (at most weakly on interspaces, medially; Figs 5K , 8 ). [Body length 3.4–6.1 mm]....................................................................................... S. (Astomyrme) pusillus (Klug in Waltl, 1835)

- Vertex with spot of golden recumbent pubescence ( Figs 13G , 15A ). T2 with lateral spots dense, well defined ( Figs 13J , 14 ). Pronotal anterior margin black ( Figs 13K , 15B ). [Body length 6.2–7.0 mm]..... S. (Astomyrme) matritensis ( Mercet, 1905)

3. T5 with full band of pale pubescence. [Scutellar scale large and elongate (ca. 2 × broader than long), shaped as a dorsally arcuate trapezium, with well formed convergent sides and apically truncate; this structure is approached only by some specimens of S. sulcisius Invrea ]................................................ S. ceballosi Suárez, 1959 (species incertae sedis)

- T5 with mostly black pubescence. [Scutellar scale different, with no elongated sides formed (if scutellar scale similar, metasomal pattern different)]..................................................................................... 4

4. Pronotum not darkened (exceptionally, weakly darker brownish anteriorly, never black). [T2 without basolateral spots of pale pubescence (only sparse setae present at most)]............................................................. 5

- Pronotum with black transverse band. [T2 with or without basolateral spots of pale pubescence]...................... 6

5. Pygidium with striae subparallel to the longitudinal axis. Scutellar scale relatively broad (e.g. ca. one sixth of mesosoma width; Petersen 1988). Dorsum of mesosoma sub-horizontal (lateral view). T2 with apical band of pale pubescence uniformly narrow. [Vertex with or without pale recumbent pubescence].......................................................................................... ‘complex S. (Smicromyrme) rufipes ’ [sensu Schmid-Egger & Schmidt 2021, 2022] 1

- Pygidium with divergent striae. Scutellar scale relatively narrow (e.g. ca. one eighth of mesosoma width; Petersen 1988). Dorsum of mesosoma angled towards scutellar scale (lateral view). T2 with apical band of pale pubescence medially broader. [Vertex with pale recumbent pubescence]............ S. (Smicromyrme) ruficollis ( Fabricius, 1793) [sensu Petersen 1988]

6. Mandible with distal element externally straight and elongate; basal groove and inner basal ridge approximately straight; basal portion very short. Dorsal mid-line approximately straight, slanted towards scutellar scale (lateral view). [Mesosoma elongate, trapezoidal. T2 with medially enlarged apical band and T3 with full band of pale pubescence, both typically continuous. T2 laterally with or without well defined spots of pale pubescence]................. .. S. (Eremotilla) sulcisius Invrea, 1955

- Mandible with distal element distally bent, convex; basal groove and inner basal ridge notably curved inwards; basal portion proportionately long. Dorsal mid-line weakly convex, with dorsum sub-horizontal (lateral view). [Band of T3 apicomedially weakly narrower, or partially to fully interrupted]........................................................... 7

7. T2 with medially enlarged apical band and T3 with full band of pale pubescence, both continuous. T2 laterally without or with weakly defined spots of pale pubescence. Scutellar scale narrow, small and inconspicuous. Clypeus with subapical tubercle on each anterolateral corner of lower half. [Pygidium basally widened, oval. Mesosoma subtrapezoidal. Mandible robust, not elongate, with distal element convex]........................ S. plantourianus Schwartz, 1986 (species incertae sedis)

- T2 basolaterally with two well defined oval spots of pale pubescence, one on each side (always visible in dorsal view). Clypeus without subapical tubercles on lower half. Scutellar scale not large, but well developed. [Pygidium elongate, may be basally widened and oval. Mandible more slender, more elongate].................................................... 8

8. T2 with medially enlarged continuous broad apical band and T3 with full band of pale pubescence apicomedially variably narrowed. Pygidium typically subtriangular........................... S. suarezi Matias, 2024 (species incertae sedis)

- T2 with apical band and T3 with band of pale pubescence, both medially fully interrupted by black pubescence. Pygidium typically elongate, subparallel sided............... S. partitus (Klug in Waltl, 1835) and S. fasciaticollis ( Spinola, 1843) [females of both species currently not clearly distinguishable] (species incertae sedis)

MALES (unknown for S. ceballosi Suárez, 1959 and S. suarezi Matias, 2024 )

1. Mandible externo-basally with no large tooth (e.g. Fig. 4F ).................................................... 2

- Mandible externo-basally with large tooth.................................................................. 4

2. Metasoma with pale pubescence forming sparse fringes only (e.g. Fig. 19B ). Radial vein with notable thickening ( Fig. 4H ).. ................................................................................................... 3

- Metasoma with pale pubescence forming medially enlarged apical band on T2 and full band on T3 ( Fig. 12I ). Radial vein without notable thickening ( Fig. 12H )................................... S. (Astomyrme) matritentis ( Mercet, 1905)

3. Propodeum red or weakly darker ( Figs 19A,C )........................ .. S. (Astomyrme) pusillus (Klug in Waltl, 1835)

- Propodeum black ( Fig. 19E )........................................... S. (Astomyrme) opistomelas Invrea, 1950

4. Mandible apically bidentate............................................................................ 5

- Mandible apically tridentate........................................................................... 10

5. Clypeus with two subapical tubercles; metasoma lacking bands of pale pubescence, only apical fringes present.......... 6

- Clypeus more or less flat without tubercles, apical margin approximately straight (may be weakly sinuate or medially weakly indented); metasoma with full bands of pale pubescence over T3 or T4.......................................... 7

6. Clypeus with distinct medial longitudinal carina, connected to two subapical tubercles.............................................................. .. ‘complex S. (Smicromyrme) rufipes ’ [sensu Schmid-Egger & Schmidt 2021, 2022] 1

- Clypeus with distally reduced medial longitudinal carina, not connected to two subapical tubercles.................................................................................... S. (Smicromyrme) ruficollis ( Fabricius, 1793)

7. T4 with mostly dark pubescence......................................................................... 8

- T4 covered with band of pale pubescence.................................................................. 9

8. Radial vein with notable thickening; second transverse-cubital vein “T-shaped”. T6 and T7 covered with white pubescence. F1 longer than 2 × pedicel length. [Mesosoma ferruginous-red, except for black medial pronotal spot, post-scutellum, sternum and mesopleuron].......................................................... S. (Eremotilla) merceti ( André, 1903)

- Radial vein without thickening; second transverse-cubital vein arcuate or “L-shaped”. T6 and T7 covered with dark pubescence. F1 as long as ca. 2 × pedicel length. [Mesosoma typically ferruginous-red, except for black pronotum laterally, meso and metapleuron and sternum; mesosoma black with red propodeum on form notomelanica Suárez, 1954].............................................................................. S. (Eremotilla) metanotalis andradei Suárez, 1954

9. Clypeus with anterior margin approximately straight. Wings yellowish-brown. Tegula dark brown, densely punctate except for impunctate outer margin. Mesosoma typically all black, except for red propodeum......................................................................................... S. (Eremotilla) metanotalis metanotalis (André, 1902)

- Clypeus with weak medial concave indentation on the anterior margin. Wings hyaline or sub-hyaline. Tegula blackish, sparsely punctate. Mesosoma typically black, except for red pronotum, mesoscutum, scutellum, postscutellum, mesopleuron, metapleuron and propodeum, the mesoscutum darkened laterally................ .. S. (Eremotilla) sulcisius Invrea, 1955

10. Mandible externo-basally with small blunt tooth (may be hidden in pubescence). Clypeus without tubercles, with longitudinal punctate medial carina on posterior half. Pale pubescence forms apical fringes on T2–4. [Anterior margin of clypeus with narrowed sides, medially narrowly emarginate]...................... S. ferdinandi Invrea, 1953 (species incertae sedis)

- Mandible externo-basally with well developed tooth. Clypeus frequently with tubercles, upper surface concave. Pale pubescence forms apical fringe on T2 and full bands on T3–4........................................................... 11

11. Clypeus with anterior margin having two forward-facing tubercles separated by medial concave depression. Apical band of T2 narrow, frequently anteromedially emarginated............................................................ 12

- Clypeus with anterior margin weakly upturned forming divergent tubercle on each anterolateral corner, both medially separated by weakly concave notch. Apical band of T2 broad, medially as broad as bands of T3–4.................................................................................... S. plantourianus Schwartz, 1986 (species incertae sedis)

12. Mesoscutum dark brownish-red to ferruginous-red, typically concolorous with pronotum, scutellum, metanotum and propodeum (some areas may be slightly darkened) [Wings strongly infuscate. Tibiae and tarsi with dark pubescence]........................................................... S. partitus (Klug in Waltl, 1835) [typical form] (species incertae sedis)

- Mesoscutum black................................................................................... 13

13. Mesoscutum strongly contrasting with orange-red pronotum, scutellum, metanotum and propodeum (mesoscutum orange-red along the pronotal suture). [Wings slightly infuscate. Tibiae and tarsi with pale pubescence]................................................................................... S. fasciaticollis ( Spinola, 1843) (species incertae sedis)

- Mesoscutum and pronotum black ........................................................................ 14

14. Mesosoma all black, except for red propodeum................................ S. partitus propodealis Suárez , 19592

- Mesosoma all black................................................... S. partitus obscurithorax (André, 1902) 2

Remarks on the key.

1 Smicromyrme rufipes dusmeti ( Mercet, 1905) , described from Sierra de Albarracín (Teruel, Spain), is not included in the key due to unresolved issues; further to the notes in Matias 2023b, this taxon was synonymized with S. verhoeffi Suárez, 1959 , by Mingo & Compte (1963: 91), which in turn would represent a new species for the Iberian Peninsula (hitherto known from Southern France); however, note the name dusmeti would have precedence over verhoeffi , which would become its junior synonym; Art. 23 of the Code (ICZN 1999); the species name would thus become S. dusmeti ( Mercet, 1905) . [Additional data on this taxon is presented by Parejo-Pulido & Romano in press].

2 Smicromyrme partitus propodealis and S. partitus obscurithorax may constitute chromatic variation of S. partitus and not true subspecies in the biological sense.

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 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 12. Smicromyrme matritensis (Mercet), male, Ribeira do Torgal (Odemira, Beja, Portugal), 15.vii.2019 (RMC817). 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 metasoma (T1–T3), dorsal view J metasoma (T2, S2; felt-lines), lateral view K genitalia, dorsal view L genitalia, ventral view M genitalia, lateral view. Scale bar: 1 mm (A–B, G), 0.25 mm (C, I–J), 0.1 mm (D), 0.2 mm (E–F, K–M), 0.5 mm (H).

FIGURE 13. Smicromyrme matritensis (Mercet), female, Ribeira do Torgal, 15.vii.2019 (RMC818). A dorsal habitus B ventral habitus C lateral habitus D pygidium E antenna F mandible, ventral view G head, frontal view H mesosoma, dorsolateral view I clypeus (white arrows indicate anterolateral tubercles) J metasoma, felt-line, lateral view K mesosoma, dorsal view L scutellar scale (white arrow). Scale bar: 1 mm (A–C), 0.1 mm (D, I, L), 0.5 mm (E, J), 0.2 mm (F), 0.25 mm (G–H).

FIGURE 14. Smicromyrme matritensis (Mercet), female, dorsal habitus, variation.A Pitões das Júnias, 24.v.2017 (RMC499) B Lamosa, 11.vi.2017 (RMC529) C Galicia, 14.v.2019 (RMC678). Scale bar: 1 mm (A–C).

FIGURE 15. Smicromyrme matritensis (Mercet), female, Galicia (Spain), 14.v.2019 (RMC678), variation. A head, frontal view B mesosoma. Scale bar: 0.5 mm.

FIGURE 18. Smicromyrme opistomelas Invrea, male, Almería (Spain), 12.vii.2023 (RMC2092). 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), 0.25 mm (C–F), 0.5 mm (H), 0.1 mm (I–K).

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).