Traumatomutilla quadrinotata (Klug, 1821)

Bartholomay, Pedro R., Williams, Kevin A., Cambra, Roberto A., Lopez, Vinícius M. & Oliveira, Marcio L., 2025, Revision of the Traumatomutilla quadrinotata species-group (Hymenoptera: Mutillidae, Sphaeropthalminae): new synonyms, sex associations, and a new species from the Brazilian Atlantic Forest, European Journal of Taxonomy 995, pp. 1-75 : 5-10

Traumatomutilla quadrinotata species-group

Diagnosis

Female

Lateral margins of the mesonotum projected laterally into blunt tubercles; head unarmed on posterior margin of vertex; scutellar scale narrow, usually lacking anterolateral carinae; pygidial plate narrow, sides strongly convergent basad, subpyriform.

Male

Mesopleuron always tuberculate on dorsal half; axillar projections always truncate; S2 always lacking setae filled pit; hypoygium elongate, subrectangular, never defined by lateral carinae; cuspis slender, elongate, mostly asetose; paracuspis poorly developed, lobe-like.

Included taxa

Traumatomutilla ameliae Casal, 1969 , T. austera ( Gerstaecker, 1874) , T. chrysozona ( Gerstaecker, 1874) , T. funebris ( Gerstaecker, 1874) , T. incerta ( Spinola, 1841) , T. infernalis ( Gerstaecker, 1874) , T. pompiliformis ( Gerstaecker, 1874) , T. quadrinotata ( Klug, 1821) , T. quadripustulata ( Klug, 1821) , T. sancta ( Gerstaecker, 1874) , T. tetratrauma Bartholomay & Williams sp. nov., and T. ursina ( Gerstaecker, 1874) .

Distribution

Widely distributed in South America from Colombia to Argentina, except Chile.

Remarks

The T. quadrinotata species-group is a more or less consistently large-bodied (> 10 mm) group in both sexes. The anterolateral carinae in the scutellar area – albeit partially concealed by dense setation – and the dorsolaterally expanded occipital carina observed in T. tetratrauma sp. nov. were not included in the original diagnosis of the T. quadrinotata species-group by Williams et al. (2017). Apart from these two new characters, the remaining characters for the species-group are relatively consistent and nonvariable for both sexes, most notably so for males, which apart from the length of certain genitalia setae and sculpture of the pronotal dorsum, can only be distinguished based on color and setae characters. The most conspicuous and reliable character for identifying females is the lateral expansion of the mesonotum which varies slightly from broad and blunt (e.g., T. funebris and T. quadrinotata ) to narrow and tuberculiform (e.g., T. ameliae and T. sancta ). The post-mesonotal tubercle is a character that was first observed in T. poranga Bartholomay & Williams, 2018 and, though it can be found in four species of the T. quadrinotata species-group, it is most conspicuous in T. quadripustulata .

Key to females of the T. quadrinotata species-group

1. Lateral surface of metapleuron and propodeum almost entirely concealed by dense golden appressed setae ( Figs 12B , 18B ); sculpture sparsely foveolate-punctate with densely micropunctate intervals where visible; vertex always clothed with black setae ( Figs 12A , 18A ) (mostly restricted to the Atlantic Forest domain) .................................................................................................................... 2

– Lateral face of metapleuron and propodeum generally smoother, at most with sparse erect silvery-white setae (e.g., Fig. 7C ); sculpture entirely exposed, sparsely foveolate-punctate with smooth, shining unsculptured intervals (e.g., Figs 1C , 8B , 14B ); vertex sometimes clothed with silvery-white setae (e.g., Fig. 8D ) ........................................................................................................................... 3

2. Frons with patch of golden setae ( Fig. 18B ); occipital carina slighty swollen dorsolaterally; anterolateral carinae present in scutellar area; dorsal mesosomal setae, when present, more or less uniform in length ( Fig. 18B ) ................................. T. tetratrauma Bartholomay & Williams sp. nov.

– Head setae entirely black ( Fig. 12A–B ); occipital carina equally wide throughout; anterolateral carinae absent in scutellar area; pronotum and anterior portion of mesonotum with numerous long erect setae when present ( Fig. 12B ) ..................................................... T. quadrinotata ( Klug, 1821)

3. T2 marked with two round orange to reddish integumental spots ( Fig. 8A, D, F, H ); mesonotum with transverse silvery band of setae, sometimes largely obliterated ( Fig. 8A, D, F, H ); occipital carina slightly swollen dorsolaterally (commonly found in the northern Amazon) ........................... .................................................................................................................... T. incerta ( Spinola, 1841)

– T2 generally with four yellow to red integumental spots (e.g., Figs 1A , 7A , 14A ), anterior spots sometimes reduced or obliterated, posterior spots sometimes enlarged and confluent; mesonotum setae generally either entirely black (e.g., Fig 14A ) or with lateral longitudinal stripes of silver to golden setae (e.g., Fig. 17A, D ); occipital carina equally wide throughout ..................................... 4

4. Head and mesosomal dorsum with setae entirely black, silvery-white setae greatly reduced on entire body, often completely absent ( Figs 4A–B , 14A–B ) ........................................................................ 5

– Silvery-white setae always conspicusouly present at least on propodeal dorsum and/or vertex (e.g., Fig. 3E ) ............................................................................................................................................. 6

5. Post-mesonotal tubercle present, sometimes reduced but still conspicuous; lateral surface of propodeum usually with large, unsculptured smooth area posteriorly; silvery-white setae present, though greatly reduced, on different areas of T2–4, S1–4, and often on metapleuron ( Fig. 14B ) (mostly found in the southern Amazon) ......................................... T. quadripustulata ( Klug, 1821)

– Post-mesonotal tubercle absent; lateral surface of propodeum usually uniformly and densely areaolatepunctate to foveolate-punctate; body setae almost all black( Fig. 4A–B ); if different color setae present, then of a coppery-golden tone and reduced to inconspicuous patches on fringes of T2–4 medially and/or laterally ( Fig. 5B ) (mostly found in Cerrado areas) ......... T. chrysozona ( Gerstaecker, 1874)

6. T2 virtually devoid of sculpture anterolaterally, smooth, shining and asetose ( Fig. 20A, C ); in addition to dense short black and silvery-white setae, entire body (except mandibles, antennae, and tarsi) clothed with dense conspicuously long brownish setae (known only from Cerrado areas so far) ( Fig. 20A, C ) ........................................................................................ T. ursina ( Gerstaecker, 1874)

– T2 at most with sparser sculpture and setation anterolaterally; body setae usually of equal length throughout, either black or silvery-white, never brownish (e.g., Fig. 1A–C ) .................................. 7

7. Post-mesonotal tubercle absent, at most with slight swelling of lateral margin posterior to mesonotal projection; T2 with orange integumental spots which can sometimes be dulled in color (found only in Cerrado/Atlantic Forest transition zones so far) ( Fig. 7A–C ) ...... T. funebris ( Gerstaecker, 1874)

– Post-mesonotal tubercle present, often reduced but still conspicuous; T2 integumental spots yellowish or reddish (e.g., Figs 3A , 17A ) ......................................................................................... 8

8. Integumental spots of T2 with scatterd foveolations usually restricted to the edges ( Fig. 17A ); spots yellowish in color, often linear in shape (commonly found in Cerrado and Caatinga areas) ( Fig. 17D ) ............................................................................................ T. sancta ( Gerstaecker, 1874)

– Integumental spots of T2 with sparse foveolations throughout; spots reddish, always subquadrate or subrectangular (e.g., Figs 1A , 3E ) .................................................................................................... 9

9. Lateral surface of propodeum densely areolate-punctate with smooth flat intervals; intervals conspicuously broader along posterior margin of lateral propodeal surface; vertex and mesonotum conspicuously clothed with silvery-white to silvery-golden appressed setae (known from Cerrado, Pampa and Atlantic Forest) ( Fig. 3A, E ) ............................................ T. austera ( Gerstaecker, 1874)

– Lateral surface of propodeum densely areolate-punctate with sharp homogeneous intervals ( Fig. 1C ); vertex and mesonotum clothed with black setae only (common in Chaco, Pampa, and xeric regions of Argentina) ( Fig. 1A ) ................................................................................... T. ameliae Casal, 1969

Key to the known color forms of the T. quadrinotata species-group females

1. Mesonotum with transverse band of silvery or golden setae ............................................................ 2

– Mesonotum entirely black, with parallel longitudinal stripes of silvery setae, or with separated spots of silvery setae .................................................................................................................................. 7

2. T2 having four integumental spots ................................................................................................... 3

– T2 having two integumental spots .................................................................................................... 4

3. Head clothed entirely with black setae ............................................................................................... ....................................... T. quadrinotata ( Klug, 1821) [nominotypical and only known color form]

– Frons clothed with golden setae ......................................................................................................... ........ T. tetratrauma Bartholomay & Williams sp. nov. [nominotypical and only known color form]

4. Spots of T2 broadly confluent ........................................................................................................... 5

– Spots of T2 widely separated ............................................................................................................ 6

5. Vertex clothed with silvery-white setae .............................................................................................. ......................................... T. incerta ( Spinola, 1841) [color form found in Peru, Huanuco province]

– Vertex clothed with black setae .......................................................................................................... .. T. incerta ( Spinola, 1841) [color form found in southwestern Brazilian Amazon, Rondônia State]

6. Vertex clothed with silvery-white setae .............................................................................................. ...................................... T. incerta ( Spinola, 1841) [formerly known as T. weyrauchi Mickel, 1952 ]

– Vertex clothed with black setae ............................. T. incerta ( Spinola, 1841) [nominotypical form]

7. Mesosomal dorsum clothed entirely with black setae ...................................................................... 8

– Mesosomal dorsum having longitudinal white stripes at least on propodeum ................................. 9

8. Spots of T2 yellow; pleural setae white; T2–5 having white setae spots medially ............................ ...................................................................... T. quadripustulata ( Klug, 1821) [nominotypical form]

– Spots of T2 red; pleural setae black; T2–5 clothed entirely with black setae ..................................... ................ T. chrysozona ( Gerstaecker, 1874) [formerly known as T. lugubrina ( Dalla Torre, 1897) ]

9. Mesosomal stripes of silvery-white setae restricted to propodeum ................................................ 10

– Mesosomal stripes, even if absent from propodeum, extending at least onto mesonotum .............11

10. Tibial spurs white ................. T. ameliae ( Casal, 1969) [nominotypical and only known color form]

– Tibial spurs black ...... T. funebris ( Gerstaecker, 1874) [nominotypical and only known color form]

11. Mesosomal stripes divided, mesonotum with broad spots of silvery-white setae; T2 with red integumental spots .......................................................................................................................... 12

– Meosomal stripes continuous; T2 with yellow integumental spots ................................................ 13

12. Vertex with medial spots of silvery-white setae ................................................................................. ......................................................................... T. austera ( Gerstaecker, 1874) [nominotypical form]

– Vertex clothed with black setae only .................................................................................................. ........................ T. austera ( Gerstaecker, 1874) [formerly known as T. sigillata ( Gerstaecker, 1874) ]

13. Vertex with transverse band of silvery-white setae ............................................................................ ....... T. sancta ( Gerstaecker, 1874) [color form found in Bolivia, Paraguay, and Midwestern Brazil]

– Vertex clothed entirely with black setae ......................................................................................... 14

14. Stripes of mesosoma extending to pronotum; with long erect setae throughout body dorsum .......................... T. ursina ( Gerstaecker, 1874) [nominotypical and only known color form]

– Stripes of mesosoma terminating on mesonotum; all dorsal setae short and mostly appressed ..... 15

15. Posterior integumental spots of T2 ovate or subquadrate, widely separated ...................................... ........................................................................... T. sancta ( Gerstaecker, 1874) [nominotypical form]

– Posterior integumental spots of T2 transversely linear, narrowly separated ...................................... ............................... T. sancta ( Gerstaecker, 1874) [formerly known as T. solemnis ( Cresson, 1902) ]

Key to males of the T. quadrinotata species-group

1. Apex of cuspis with short setae ( Figs 9E , 19E ) ................................................................................ 2

– Apex of cuspis with long setae ......................................................................................................... 3

2. Pronotum sparsely and finely foveolate-punctate with micropunctate intervals, concealed by dense silvery-white setation ( Fig. 9A ) (predominantly found in the Amazon Forest areas) ........................ ..................................................................................................................... T. incerta ( Spinola 1841)

– Pronotum densely and finely foveolate-punctate without micropunctured intervals, sculpture mostly exposed, setae black ( Fig. 19A ) (predominantly found in Atlantic Forest areas) .............................. ............................................................................... T. tetratrauma Bartholomay & Williams sp. nov.

3. Lateral suface of propodeum and metapleuron clothed with dense appressed silvery-golden setae .. .............................................................................................................. T. quadrinotata ( Klug, 1821)

– Lateral surface of propodeum at most with sparse erect silvery-white setae; metapleuron at most with sparse appressed black or silvery-white setae .......................................................................... 4

4. Body setae predominantly black with coppery-golden areas ...... T. chrysozona ( Gerstaecker, 1874)

– Body setae predominantly black with silvery-white areas ............................................................... 5

5. Wings brown infuscated on apical third, hyaline-brown elsewhere; T2 clothed entirely with silvery-white setae ...................................................................................... T. quadripustulata ( Klug, 1821)

– Wings entirely dark brown with strong violaceous and blueish reflections; T2 almost entirely clothed with black setae or at most with silvery-white setae on anterior half ............................................... 6

6. Silvery-white setae markings of propodeum and T2 dense, completely obscuring integument beneath ............................................................................................................ T. ameliae Casal, 1969

– Silvery-white setae markings of propodeum and T2 sparse, if apparently dense, then never completely obscuring integument beneath .......................................................................................................... 7

7. Cuspis length approximately 0.65 × paramere length; paracuspis relatively well developed, almost node-like ( Fig. 11F, N ); posterior margin of penis valve subtruncate; silvery-white setae present throughout propodeal dorsum .................................................. T. pompiliformis ( Gerstaecker, 1874)

– Cuspis length approximately 0.85 × paramere length; paracuspis poorly developed, lobe-like; posterior margin of penis valve evenly convex; silvery-white setae restricted to posterior half of propodeal dorsum ........................................................................... T. infernalis ( Gerstaecker, 1874)

Key to the known color forms of the T. quadrinotata species-group males

1. Body with black and golden to cupreous setae ................................................................................. 2

– Body with black and silvery-white setae .......................................................................................... 5

2. Wings with basal third hyaline-brown, conspicuously less infuscated than remainder of wing membrane ..................... T. quadrinotata ( Klug, 1821) [nominotypical and only known color form]

– Wings completely dark brown, without any conspicuously lighter areas ........................................ 3

3. Lateral surface of propodeum clothed with golden setae only ........................................................... ........ T. tetratrauma Bartholomay & Williams sp. nov. [nominotypical and only known color form]

– Lateral surface of propodeum clothed with black setae only ........................................................... 4

4. Propodeum and base of T2 clothed with dense golden setae ............................................................. ............................... T. chrysozona ( Gerstaecker, 1874) [formerly known as T. dives ( André, 1906) ]

– Propodeum and base of T2 clothed having mostly black setae, at most having some scattered golden setae .......................................................... T. chrysozona ( Gerstaecker, 1874) [nominotypical form]

5. Pronotum densely clothed with silvery-white setae ......................................................................... 6

– Pronotum clothed predominantly with black setae, if silvery-white setae present, then always sparse and mostly erect ................................................................................................................................ 9

6. Fringe of T2–3 clothed only with black setae medially ...................................................................... ................................................. T. ameliae Casal, 1969 [nominotypical and only known color form]

– Fringe of T2–3 clothed with silvery-white setae throughout ............................................................ 7

7. Fringe of T4 clothed with silvery-white setae throughout .................................................................. ............................ T. quadripustulata ( Klug, 1821) [formerly known as T. maraca ( Cresson, 1902) ]

– Fringe of T4 clothed with black setae medially and silver-white setae laterally .............................. 8

8. S4 fringe with black setae ................................................................................................................... .......................................... T. incerta ( Spinola, 1841) [formerly known as T. dentata ( Smith, 1879) ]

– S4 fringe with silvery-white setae .......... T. incerta [formerly known as T. dignitosa Mickel, 1952 ]

9. T2 clothed almost entirely with silvery-white setae, more densely on anterior third ......................... ....................... T. quadripustulata ( Klug, 1821) [formerly known as Mutilla pruinosa Smith, 1855 ]

– T2 clothed predominantly witth black setae, at most with patches of silvery-white setae on anterior third ................................................................................................................................................. 10

10. Propodeal setae mostly black, silvery-white setae restricted to posterior surface of propodeum ...... ................................................. T. infernalis ( Gerstaecker, 1874) [nominotypical form and formerly known as T. floccos ( Gerstaecker, 1874) ,no discernible difference between holotypes]

– Silvery-white setae present throughout propodeal dorsum ........ T. pompiliformis ( Gerstaecker, 1874) [nominotypical form and formerly known as T. serra ( Cresson, 1902) , no discernible difference between holotypes]

Figures

Fig. 12. Traumatomutilla quadrinotata (Klug, 1821), lectotype, ♀ (ZMB). A. Dorsal habitus. B. Lateral habitus. C. Type labels. Scale bars = 2 mm.

Fig. 18. Traumatomutilla tetratrauma Bartholomay & Williams sp. nov., holotype, ♀ (MZSP). A. Dorsal habitus. B. Lateral habitus. Scale bars = 2 mm.

Fig. 7. Traumatomutilla funebris (Gerstaecker, 1874), holotype, ♀ (ZMB). A. Dorsal habitus. B. Type labels. C. Lateral habitus. Scale bars = 2 mm.

Fig. 1. Traumatomutilla ameliae Casal, 1969, holotype, ♀ (AMNH). A. Dorsal habitus. B. Type labels. C. Lateral habitus. Scale bars = 5 mm.

Fig. 8. A–C. Traumatomutilla sodalicia (Kohl, 1882), holotype, ♀ (NHMW). A. Dorsal habitus. B. Lateral habitus. C. Type labels. D–E. Traumatomutilla tabatinga Casal, 1969, holotype, ♀ (AMNH). D. Dorsal habitus.E. Type labels.F–G. Traumatomutilla incerta (Spinola, 1841), ♀ (non-type specimen), most common color form. F. Dorsal habitus. G. Lateral habitus. H. Traumatomutilla incerta (Spinola, 1841), ♀ (non-type specimen), color form with reduced silvery-white setae on head and mesosoma, dorsal habitus. Scale bars: A–B = 5 mm; D, F–H = 2 mm.

Fig. 14. Traumatomutilla quadripustulata (Klug, 1821), lectotype, ♀ (ZMB). A. Dorsal habitus.B. Type labels. C. Lateral habitus. Scale bars = 2 mm.

Fig. 17. A–C. Traumatomutilla sancta (Gerstaecker, 1874), holotype, ♀ (ZMB). A. Dorsal habitus. B. Lateral habitus. C. Type labels. D–F. Traumatomutilla solemnis (Cresson, 1902), holotype, ♀ (CM). D. Dorsal habiuts. E. Lateral habitus. F. Type labels. Scale bars = 2 mm.

Fig. 4. Traumatomutilla chrysozona (Gerstaecker, 1874), ♀ (non-type specimen). A. Dorsal habitus. B. Lateral habitus. Scale bars = 5 mm.

Fig. 3. A–C. Traumatomutilla austera (Gerstaecker, 1874), lectotype, ♀ (ZMB). A. Dorsal habitus. B. Lateral habitus. C. Type labels. D–F. Traumatomutilla sigillata (Gerstaecker, 1874), holotype, ♀ (ZMB). D. Dorsal habitus. E. Lateral habitus. F. Type labels. Scale bars = 2 mm.

Fig. 5. Traumatomutilla chrysozona (Gerstaecker, 1874), lectotype, ♂ (ZMB). A. Lateral habitus. B. Vestigial coppery setae on female metasoma, lateral view. C. Type labels. D. Genitalia (halved), dorsal view. E. Genitalia (halved), ventral view. F. Genitalia (halved, penis valve removed), lateral/ inner view. G. Cuspis (removed, not to scale), lateral/inner view. H. Penis valve (removed, not to scale), lateral/outer view. Scale bar = 2 mm.

Fig. 20. Traumatomutilla ursina (Gerstaecker, 1874), holotype, ♀ (ZMB). A. Dorsal habitus. B. Type labels. C. Lateral habitus. Scale bars = 2 mm.

Fig. 9. Traumatomutilla dentata (Smith, 1879), holotype, ♂ (NHMUK). A. Lateral habitus. B. Genitalia (halved), dorsal view. C. Genitalia (halved), ventral view. D. Genitalia (halved, penis valve removed), lateral/inner view. E. Cuspis (removed, not to scale), lateral/inner view. F. Penis valve (removed, not to scale), lateral/outer view. G. Type labels. Scale bar = 2 mm.

Fig. 19. Traumatomutilla tetratrauma Bartholomay & Williams sp. nov., allotype, ♂ (MZSP). A. Lateral habitus. B. Genitalia (halved), dorsal view. C. Genitalia, (halved), ventral view. D. Genitalia (halved, penis valve removed), lateral/inner view. E. Cuspis (removed, not to scale), lateral/inner view. F. Penis valve (removed, not to scale), lateral/outer view. Scale bar = 2 mm.

Fig. 11.A–H.Traumatomutilla pompiliformis(Gerstaecker, 1874),lectotype, ♂(ZMB).A. Dorsalhabitus. B. Lateral habitus. C. Genitalia (halved), dorsal view. D. Genitalia, (halved), ventral view. E. Genitalia (halved, penis valve removed), lateral/inner view. F. Cuspis (removed, not to scale), lateral/inner view. G. Penis valve (removed, not to scale), lateral/outer view. H. Type labels. I–P. Traumatomutilla serra (Cresson, 1902), holotype, ♂ (CM). I. Dorsal habitus. J. Lateral habitus. K. Genitalia (halved), dorsal view. L. Genitalia, (halved), ventral view. M. Genitalia (halved, penis valve removed), lateral/inner view. N. Cuspis (removed, not to scale), lateral/inner view. O. Penis valve (removed, not to scale), lateral/outer view. P. Type labels. Scale bars: A–B = 5 mm; I–J = 2 mm.