Revision of the Traumatomutilla gemella species-group (Hymenoptera, Mutillidae) with the description of its hitherto unknown males

ABSTRACT Traumatomutilla André, 1901 is a genus of Mutillidae Latreille, 1802 with 14 species-groups, of which the T. gemella group is one of the most poorly known. The hitherto unknown males of Traumatomutilla diophthalma (Klug, 1821), T. chuza Casal, 1969, and T. gemella André, 1906 are described and illustrated. The females of T. andrei (Cresson, 1902), T. angustata (André, 1906), T. chuza Casal, 1969, T. diopthalma (Klug, 1821), and T. gemella (André, 1906) are redescribed. A new species, T. peismatara Bartholomay & Cambra, n. sp. (female from Peru, male from Brazil) is described. Two host records (Podium sp. from Sphecidae Latreille, 1802 and Trypoxylon sp. from Crabronidae Latreille, 1802) are provided for T. diophthalma based on trap nest data. Traumatomutilla rastra Casal, 1969 is recognized as a new synonym of T. angustata (André, 1906).


INTroDUCTIoN
Velvet ants (Mutillidae Latreille, 1802) are solitary wasps whose larvae act as parasitoids of encapsulated immatures of other insects, especially other solitary Hymenoptera (Brothers 2006). Females of this family are always wingless while the males are usually fully winged, rarely brachypterous or apterous (Brothers 2006). This remarkable sexual dimorphism has resulted in many species being described based only on males or females and usually differentiated solely on color and setae characters (Gerstaecker 1874;Cresson 1902;Casal 1969). The reliability of such characters alone for differentiating species in Traumatomutilla André, 1901 was brought into question by research on the closely related Dasymutilla Ashmead, 1899, which demonstrated the existence of Müllerian mimicry complexes within these wasps (Williams et al. 2011a(Williams et al. , 2012Wilson et al. 2012Wilson et al. , 2013. The influence of such mimicry complexes in the coloration of these wasps is such that several species, once thought to be distinct, have been found to belong to a single widespread and highly variable species (Williams et al. 2011a(Williams et al. , 2012. Similar results have already been found in recent revisions of other Traumatomutilla species-groups (Bartholomay et al. 2019a, b). These revisions were facilitated by Williams et al. (2017), who organized the 135 species of Traumatomutilla based on females into 14 species-groups based on shared structural characters or shared combinations of structural characters. With only six known species, the Traumatomutilla gemella species-group is one of the smallest, rarest, and most morphologically distinct species-groups within Traumatomutilla (Williams et al. 2017). In this study we review and provide the first sex associations for the T. gemella species-group, as well as two novel host records for the genus.

MATerIAL AND MeTHoDS
In the current work, we studied approximatively 100 specimens, which belong to the institutions listed below.
Under the description and redescription sections, because of the highly variable nature of the coloration and setal patterns in Traumatomutilla, we provide a separate section titled "Coloration and variations", in which we provide an overall description of the known color and setal patterns for a given species.
In the Material examined section, abbreviations, acronyms, and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of each Material examined section.
AbbreviAtions Morphology T1-6 (in females), T1-8 (in males): metasomal tergites 1 to 6 and 1 to 8; S1-6 (in females), S1-8 (in males): metasomal sternites 1 to 6 and 1 to 8; IoD interocellar distance (males only); ooD ocellar-ocular distance (males only); DLo diameter of lateral ocellus (males only). Traumatomutilla gemella species-group diAgnosis. -Females. Females of this species-group can be distinguished by a single autapomorphy: the pygidial plate with projected flange-like lateral carina restricted to apical fourth. The following combination of characters is unique to the T. gemella group females, though some characters may occur in other groups as well: body generally slender, elongate; head unarmed posterolaterally; clypeus shallowly but conspicuously bilobate on apical/ventral margin, longitudinally elevated medially; vertex and/or front frequently with medial longitudinal carina; pronotal collar with vestigial transverse rugosities anteriorly; anterior face of pronotum short, shorter than or as long as pronotal collar; lateral face of pronotum with subacute tubercle anteroventral in relation to pronotal spiracle; pronotal and propodeal spiracles almost flat against lateral margin of mesosoma; mesosoma almost straight laterally, at most slightly divergent anterad, not constricted anterior to propodeal spiracle; dorsal face of propodeum much longer than posterior face; scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area; and apex of middle and hind femora rounded. Males (hitherto unknown). The males can be distinguished by a unique genitalic autapomorphy: the cuspis is at most 0.6 × the length of the paramere, abruptly upcurved at midlength, with long twisted setae along the ventral surface of the apical half. The following combination of characters is unique to the T. gemella group males, though some characters may occur in other groups as well: integument black to reddish-black with contrasting black and silvery-white to silvery-golden setae patterns varying in density; head transversely subrectangular in dorsal view; parapsis and notaulus vestigial, restricted to posterior margin of mesoscutum; axilla pronounced as twisted oblique and acute projection; scutellum gibbose, without posterior transverse carina, frequently with weak medial longitudinal carina anteriorly; mesopleuron simply swollen on dorsal half, without any projections or tubercles; meso-and metafemora rounded apically; S2 without setae-filled pit; pygidial plate strongly concave apically, apical margin strongly deflected upward; hypopygium rectangular, longer than broad, latero-apical corners angulate and slightly projected. reMArks At first glance, females of the T. gemella species-group are similar to those of the Dasymutilla paradoxa (Gerstaecker, 1874) species-group, which was recently revised by Luz et al. (2016). They differ in the presence of flange-like projections apicolaterally on the pygidial plate, absent in the D. paradoxa species-group, and the sub-petiolate T1 shape which is globose, subcylindrical in the D. paradoxa species-group. Additionally, females of the T. gemella species-group have only one pair of integumental spots on T2 while those of the D. paradoxa species-group have two pairs. Males of the T. gemella species-group can be easily separated by their axil-lae, which are produced posteriorly as short acute projections and the all black integument of T2; this contrasts with the axillar projections connected with the lateral margins of the mesoscutellum and T2 predominantly yellowish observed in males of the D. paradoxa species-group. Among the Traumatomutilla species-groups, there are no females that can be easily confused with those of the T. gemella species-group. There are males of the T. indica group that, as with the males of the T. gemella group, have the axillar projections acute and the mesopleuron unarmed. These, however, can be readily recognized by having a setae-filled pit on S2 and/or a relatively straight cuspis that lacks long sinuous setae.

Male
Unknown.

Male
Unknown.

Coloration and variations
Female (Figs 2A, c; 3A, c; 4A, c). Integument black to brownish-black except for mandibles and antennal flagella partially reddish-brown, and T2 with a pair of large subcircular orange to reddish integumental spots. Body setae predominantly silvery-yellow to golden varying in density, except for the following areas with black to brownish-black setae varying in density: gena, malar space, and ventral half of frons; pronotal dorsum, anterior half of mesonotum, propodeal dorsum medially, and femora apicodorsally; T1 medially, disc of T2 (except over integumental spots), fringe of T2 medially, fringe of T3-4 sublaterally (medial area of silvery-golden setae on T3 sometimes greatly reduced), fringe of T5 laterallly, and S6.
Vertex with or without a medial spot of silvery-yellow setae.

Male. Unknown.
reMArk Traumatomutilla angustata specimens have a single inconspicuous difference in relation to specimens of T. diophthalma which is the propodeum somewhat simply sloping posterad in lateral view (Figs 2C; 3C; 4C) as opposed to the more angulate propodeum of T. diophthalma (Fig. 7B). This propodeal structure is also seen in T. rastra, which differs from T. angustata by having the transverse band of silvery setae on The remaining structural features of T. angustata (including T. rastra n. syn.), are identical to those of T. diophthalma.
Since the few differences between T. angustata and T. rastra are restricted to minor color and setae characters which are highly variable within Traumatomutilla, we propose T. rastra as a junior synonym of T. angustata. wings (Fig. 6A). Fore wing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells; membrane dark brown, slightly but conspicuously lighter on basal third. legs (Fig. 6A). Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins.

Traumatomutilla chuza
Metasoma (Fig. 6A). ratios of width of T1, width of T2 and length of T2 45:76:58. Dorsal metasomal sculpture partially concealed by dense setation, sparsely and coarsely punctate with interspersed micropunctures where visible; pygidial plate somewhat concave, posterior margin conspicuously curved upward; surface predominantly smooth, shining, with vestigial undefined sculpture along apical margin; weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, ending in blunt, low, concave carina. S2 sparsely and finely foveolate-punctate to punctate; foveolae conspicuously sparser posterad; anteromedial crest-fold almost absent; sternal pit absent. S3 sparsely and finely punctate with sparse interspersed micropunctures; S4-7 sparsely foveolatepunctate. S7 longer than broad, with conspicuous medial longitudinal unsculptured area; posterior margin projected apicolaterally and medially; medial projection blunt, longer than lateral projections. Genitalia (Fig. 6b-F). Parapenial lobe not at all pronounced apically, simply rounded. ratios of free length of paramere, cuspis and digitus, 56:28:13; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense setae ventrally at basal half; cuspis short, stout, slightly swollen medially and narrower apicad in lateral view; narrower apicad and almost straight in dorsal view; abruptly curved dorsally in wide angle at basal third; with dense conspicuous, strongly sinuous setae on ventral surface, except at basal third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis well-developed, not sessile, slightly elongate vertically, subrounded at apical margin, densely setose along posterior margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, inconspicuously setose basodorsally; penis valve strongly concave on inner surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, with lateral pocket present on outer surface; apical distance between teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface.

Coloration and variations
Female (Fig. 5A, c). Integument black to brownish-black except for mandibles and antennal flagella partially reddishbrown, and T2 with a pair of large subcircular orange integumental spots. Body setae predominantly silvery-white except for the following areas with black setae varying in density: pronotal dorsum, anterior half of mesonotum, posterior half of mesonotum medially, propodeal dorsum medially, femora apicodorsally, T1 posteromedially, disc of T2 (except over integumental spots), fringe of T2-3 medially (except inconspicuous patch of silvery-white medially), T4 sublaterally, T5-6 (except pygidial plate) laterally, fringe of S5, and S6. Some specimens may have the head completely covered with black setae (except gena and malar space), and the silverywhite setae of the mesosomal dorsum restricted to narrow inconspicuous lateral stripes on propodeum. Such specimens have the remaining silvery-white setae areas conspicuously less defined and the integumental spots of T2 have a more reddish tone rather than the usual orange. MALe. (Fig. 6A) Integument black. Body setae predominantly silvery-white varying in density except for the following areas with black setae varying in density: mesoscutum, tegula, axillar projections, mesoscutellum, disc of T2 (except anterior third), fringe of T2-4 medially, T5-7, fringe of S5-6, and S7.  (1) T. diophthalma occurring in the same area as T. chuza (Pará, Brazil), there are no specimens of T. chuza known from West or South of Leticia (Amazonas, Colombia). Apart from the characters mentioned in the diagnosis, the females of T. chuza are easily recognizable by their more robust and stout body in comparison to the rather slender and elongate body of the other species within the T. gemella species-group. The specimen recently collected in rondônia by KAW (DZUP-299000), is undoubtedly from the T. gemella species-group and was identified as T. chuza by its overall body structure, proportions, and its occurence on the Brazilian Amazon. The coloration of this specimen, however, is remarkably different from most T. chuza since the head and mesosoma are almost devoid of silvery-white setae dorsally, except for a pair of lateral longitudinal stripes on the propodeal dorsum. This color variation may be a result of a local Müllerian mimicry pattern which includes T. barathra Williams, 2018 andT. luscoides André, 1908, species that were collected in the same area as this particular specimen. description Female body length. 13 mm.

Genitalia (Fig. 8b-F).
Parapenial lobe not at all pronounced posteriorly, simply rounded. ratios of free length of paramere, cuspis and digitus, 53:31:14; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen medially and narrower posterad in lateral view; narrower posterad and almost straight in dorsal view; abruptly curved dorsally in wide angle at anterior third; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis welldeveloped, not sessile, slightly elongate vertically, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, inconspicuously setose basodorsally; penis valve strongly concave on inner surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, with lateral pocket present on outer surface; apical distance between teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface.

Coloration and variations
Female (Fig. 7A, b). Integument black, except mandibles and antennal flagella partially reddish-brown and T2 with a pair of orange subcircular integumental spots which vary slightly in size. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: front, genae; pronotal dorsum, mesonotum medially, scutellar area, propodeal dorsum medially; T1 medially, disc of T2 (except integumental spots), fringe of T2-5 sublaterally, T6 laterally, and S6. Fig. 8A). Integument black, with mandibles and flagella partially reddish-brown. Head mostly with dense white setae except sparse, erect and large, black setae on postero-lateral areas of vertex and near inner eye margins; pronotum dorsum, mesoscutum, axillar projections, scutellum and tegula with black setae; dorsum of propodeum with sparse white setae, dense and decumbent antero-laterally; pronotum lateral face with white setae; mesopleura with sparse black setae near tegula other area with sparse white setae; propodeum lateral face with sparse white setae; legs with white setae except apex of meso and metafemora dorsally with black setae, T1 with white setae, dense and decumbent on dorsal face; T2 to T7 with black setae except anterior third and narrow lateral area of T2, narrow apical fringe of T2, narrow lateral areas of T3-4 with white setae; S1 to S4 with white setae; S5-S6 with white and black setae, S7 with black setae. reMArks This species is, to the best of our knowledge, the most widely distributed in the genus, being found from Paraguay and Mid-western Brazil to Panama. Its distribution, however, is remarkably "patchy" in between these extremes. It is not clear whether this is due to lack of sampling or its range being indeed disjunct, interrupted near the Amazon and resumed further South. The latter would be a novel distribution for Traumatomutilla, since most species seem to be widespread and common across large areas -e.g. T. ocellaris (Klug, 1821) which is found from Argentina to the southern edges of the Amazonian Forest -or restricted to certain types of environments like T. bifurca (Klug, 1821)    Metasoma (Figs 9A, c; 10A, b). ratios of width of T1, width of T2 and length of T2, 31:70:69. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolae sparser and micropunctures absent laterally and over integumental spots; sculpture less defined in general over integumental spots. T3-6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; pygidial plate broadly subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface mostly irregularly longitudinally rugose; interstice apparently granulose. S1 sparsely punctured, surface wedge-like, ending in a rounded longitudinal carina, slightly lower medially. S2 sparsely foveolate-punctate, punctures conspicuously smaller anteriorly; anteromedial crest-fold vestigial. S3-6 densely and coarsely foveolate-punctate with sparse micropunctures at S3-4; sculpture denser on S6.

Mesosoma (
Metasoma (Fig. 11A). T1 0.5 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly and vestigially rugose, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, slightly pronounced carina lower medially. S2 coarsely and sparsely foveolate-punctate to punctate, interspersed micropunctures present anterolaterally, foveolae conspicuously sparser and larger posterad; with vestigial longitudinal anteromedial crestfold; sternal pit absent. S3-5 sparsely and coarsely foveolatepunctate with interspersed micropunctures; S6-7 sparsely foveolate-punctate. S7 longer than broad, posterior margin projected laterally and medially, medial projection terminating in a pair of very small subacute closely spaced tooth-like process on posterior margin. Genitalia (Fig. 11b-F). Parapenial lobe not at all pronounced posteriorly, simply rounded. ratios of free length of paramere, cuspis and digitus, 63:33:19; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen sub-basally, narrower posterad, and almost straight in dorsal view; abruptly curved in wide angle at anterior third and narrower posterad in lateral view; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with simple short setae; paracuspis well-developed, not sessile, slightly elongate vertically, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, sparsely setose dorsally at base; penis valve strongly concave on internal surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, lateral pocket present on outer margin; distance between apex of teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface.

Coloration and variations
Female (Figs 9A, c; 10A, b). Head, mesosoma and metasoma integument always black, at most brownish-black, with ZOOSYSTEMA • 2021 • 43 (1) mandibles and antennal flagella partially reddish-brown and T2 with a pair of subrounded orange integumental spots varying in size; seta varying in density and predominantly silvery-golden except the following areas with black setae: head (except antennae), dorsum of pronotum, mesonotum, dorsum of propodeum medially, lateral face of pronotum, anterior half of mesopleuron, apex of meso and metafemora dorsally, most of disc of T2 (except integumental spots), most of fringe of T2-3 (except medially and laterally), narrow sublateral areas on fringe of T4, and S6.

reMArks
The sex association was based on distribution and matching color pattern, since both sexes of Traumatomutilla gemella are part of a well-defined mimicry syndrome occurring in the Atlantic Forest, which has been observed and used as a basis for sex associations in multiple species such as Pappognatha patruelis (André, 1898), Hoplocrates cephalotes (Swederus, 1787), Hoplomutilla spinosa (Swederus, 1787), Atlantilla auriculata (Gerstaecker, 1874), andT. guarata Casal, 1969 (KAW andPrB pers. obs.). To the best of our knowledge, the only way to differentiate the males of T. gemella from the remaining males of the species-group are the setal characters previously mentioned in the diagnosis and identification key. Setae and color characters are also the most easily observable character to differentiate females of T. gemella especially since it appears that there are no variations whatsoever in the specimens examined. In the case of the females, however, there are reliable structural characters that can be used, namely the presence of micropunctures on the lateral propodeal face.  description Female body length. 12 mm.
Genitalia (Fig. 13b-F). Parapenial lobe not at all pronounced posteriorly, simply rounded. ratios of free length of paramere, cuspis and digitus, 48:24:11; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen medially and narrower posterad in lateral view; narrower posterad and almost straight in dorsal view; abruptly curved dorsally in wide angle at anterior third; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis well-developed, not sessile, slightly elongate longitudinally, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, sparsely setose dorsally at base; penis valve strongly concave on internal surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, lateral pocket present on outer margin; distance between apex of teeth 0.1 × length of valve; dense setae present along truncate, flat posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface.

Male (
reMArks Females of T. peismatara Bartholomay & Cambra n. sp. are structurally similar to those of T. diophthalma, but differ mainly in features of the propodeum, namely the overall shape of the dorsal propodeal face and its relation with the mesonotum in lateral view. This small difference originally raised doubts about the validity of this new species and it was initially considered a variation of T. diophthalma. After the sex association of T. chuza, T. diophthalma, and T. gemella, a fourth male morphospecies remained that could not be properly placed within any of the known species, especially because of the structural differences observed in the hypopygium. These males were all collected in Amazonian areas that are relatively close to the Peruvian distribution of T. peismatara Bartholomay & Cambra n. sp. and in areas where no other females of the T. gemella species-group were found. Based on this, morphological features, and similar distribution, we hypothesize that the females of T. peismatara Bartholomay & Cambra n. sp. represent a distinct species and the males collected in the far west of the Brazilian Amazon are conspecific with those females. key to feMAles of tHe TraumaTomuTilla geMellA species-group ZOOSYSTEMA • 2021 • 43 (1)

DISCUSSIoN
The T. gemella species-group is at the same time the most widely distributed in the genus and one of the rarest, with only 100 specimens found in collections and recorded from Argentina to Panama. This disparity results in paradoxical situations; for example, T. andrei appears to be widely distributed in Brazil but is known from three specimens only. Sex associations were particularly difficult to recognize. Therefore, apart from the T. diophthalma couple reared in Panama, sex associations in this species-group relied heavily on close or overlapping distribution of males and females. This situation is not ideal, but the sex association hypotheses proposed here are supported by the morphological clues provided by the reared male specimen of T. diophthalma in Panama. Males and females of T. gemella co-occur in the Atlantic Forest and both sexes have the typical color syndrome for the region. extensive sampling over the years in the Ducke reserve, Manaus, Amazonas, Brazil, has produced only males and females of T. chuza. Additionally, males of T. chuza have not been recorded in areas outside of the Amazon so far, while T. diophthalma males have been recorded as far South as Mato Grosso state in Brazil. Finally, through a process of elimination and considering the clear differences between males of T. chuza and T. peismatara Bartholomay & Cambra n. sp. and the known distribution of T. peismatara, the sex association for this new species became obvious. Though there are obvious differences in the cuspis, digitus, and penis valve between all males of the T. gemella speciesgroup, it is still unclear how reliable and/or variable these characters are due to the small number of specimens available. At this early stage, however, it is evident that the penis valve of T. gemella is conspicuously more elongate with shorter posteroventral teeth, unlike those of the other species that are stout with a more prominent narrow anterior tooth. The differences between the genitalia of T. diophthalma and T. chuza are all superficial and can only be properly delimited with larger series of specimens.
The biological data presented here are the first records of Traumatomutilla attacking arboreal or twig-nesting hosts, albeit in trap-nests. It is important to note that females of the T. gemella species-group have the most reduced pygidial plate in the genus, with the flange-like lateral carinae restricted to the apical fifth of the plate; also, the protarsal rake is almost absent. Well developed protarsal rakes and pygidial plate have been associated with velvet-ant species that attack groundnesting hosts (Williams et al. 2011b;Bartholomay et al. 2018). The reduction of both these characters in the females of the T. gemella species-group, coupled with two different instances of them being reared from non-ground-nesting hosts, is strong evidence that this association is true. Finally, it is quite remarkable that a single species, T. diophthalma, parasitizes very distinct hosts, a cockroach-hunting sphecid (Podium) and a spider-hunting crabronid (Trypoxylon). It is important to note that the rearing of the male specimen from the Podium sp. nest was highly experimental and therefore done in very different conditions than those likely found by this species in nature. This particular host/parasite relationship, however, possibly occurs naturally since both species were collected in the same area and the female was capable of opening the chitinous cocoon of the host on its own. Additionally, Morato (1994) reared Couples of Xystromutilla asperiventris André, 1905 (Mutillidae: Sphaeropthalminae) from species of Podium and Trypoxylon in the same area.