The usage of Paraferreola Šustera, 1912 since its proposal as a genus of spider wasps, but based on a misidentified type species (Sphex ursus Fabricius, 1793, actually a species of Mutillidae), shows that it continues to be applied (although infrequently) in Pompilidae, despite the proposal of new names (Eoferreola Arnold, 1935 and Tea Pate, 1946) for the equivalent generic concept. Application of Article 70.3.2 of the fourth edition of the International Code of Zoological Nomenclature permits correction of the erroneous type species designation to that originally intended (Sphex rhombica Christ, 1791), and thus the maintenance of Paraferreola as a valid genus in Pompilidae. Examination of the holotype specimen of Sphex ursus has shown it to be a senior synonym of Mutilla vesta Cresson, 1865 and its junior synonyms, for which the valid name must thus be Dasymutilla ursus (Fabricius, 1793), comb. nov.

Dasymutilla vesta, Eoferreola, new combination, new synonymy, Tea

The family attribution of the genus Paraferreola Šustera, 1912 has long been contentious. It was originally proposed for a group of spider wasps (Pompilidae), with Sphex ursus Fabricius, 1793 designated as the type species (Šustera 1912). However, examination of the type specimen of that species by various authors showed that it is not a pompilid, but instead is a male of Mutillidae, although its genus was then not determined. Consequently, Pate (1946) excluded Paraferreola from Pompilidae and proposed the use of a different name for the relevant pompilid taxon. Nevertheless, some authors have persisted in the use of Paraferreola in Pompilidae, considering that Šustera had misidentified his type species. The name Paraferreola has never been used as a valid name in Mutillidae.

Here we review the history and usage of the names involved and propose a solution that maintains Paraferreola as an available name in Pompilidae, and identifies Sphex ursus to species in Mutillidae.

In his treatment of the Piezata (= Hymenoptera) Fabricius (1793) included 92 species in the genus Sphex Linnaeus, 1758. Many were newly described, including, on p. 210:

“Urſus. 48. S.[phex] hirta atra abdominis ſegmento ſecundo ferrugineo, alis atris.

Habitat – – Muſ. Dom. Lund. [locality unknown, specimen in collection of Niels Tønder Lund]

Statura & magnitudo praecedentis [S. viatica Linnaeus]. Corpus totum hirtum, atrum ſegmento abdominis secundo ſolo ferrugineo. Alae nigrae.”

The brief description refers to a hairy black wasp with only the second abdominal [metasomal] segment ferruginous and black wings, but from an unknown locality. At that time, Fabricius included many distantly related species in the genus Sphex. However, Fabricius (1798) established the new genus Pompilus Fabricius, 1798 to accommodate a group of species he considered very similar to Sphex itself, and included 37 species, most transferred from Sphex. The fifth species of Pompilus is P. ursus, accompanied by a description identical to the 1793 treatment, still with no indication of its place of origin. This entry was repeated in Fabricius’s (1804) work reviewing the Hymenoptera (14^th under Pompilus). The concept that S. ursus is a member of the Pompilidae thus originated with Fabricius. This was confirmed by Dalla Torre (1897: 330) in his multi-volume catalogue of the Hymenoptera, where he recognized Pompilus ursus as a valid species (from central and southern Europe), but also suggested that Sphex rhombica Christ, 1791 might be a senior synonym; he placed Pompilus coccineus Fabricius, 1804 (female) as a junior synonym and also synonymized a few other pompilid species with it. Schulz (1906: 169), in his detailed commentary on Dalla Torre’s catalogue, suggested that Pompilus amurensis Motschulsky, 1860 might actually be “ursus F.” or a relative, but made no further comment on ursus, and it is probable that he accepted Dalla Torre’s concept of the species since he did not comment on the contradiction between Motschulsky’s characterization of P. amurensis as “nitidus” (smooth) and Fabricius’s “hirta” (hairy) for S. ursus.

The first contradiction to the placement of ursus in Pompilidae was provided by Nielsen (1907) who, in his treatment of the Pompilidae of Denmark, included the species “F.[erreola] coccinea Fabr. (nec ursus Fabr.)*)” and a very brief explanatory footnote (p. 49) “*) P. ursus Fabr. er efter Typeeksemplaret en Mutilla.” implying that he had examined the type specimen and found that it was a Mutilla (used in a sense encompassing most members of the family Mutillidae, excluding Myrmosa Latreille, 1797). No further information was provided under the section on the Mutillidae, where only two species of Mutilla were included and ursus did not appear, implying that Nielsen did not consider it to be a Danish species.

Šustera (1912, 30 September) reviewed the Palaearctic genera of Pompilidae and proposed Paraferreola Šustera, 1912 as a ‘new name’ for “Ferreola Smith (Kohl, Costa, Tourn.) part., Rad., Ashm., nec. Lep[eletier de Saint-Fargeau 1845].”, and he specified “Type: Paraferreola ursus F.”. Šustera had evidently not seen Fabricius’s type material, nor Nielsen’s (1907) footnote, but merely relied on subsequent placement of ursus in Pompilus by Fabricius (1798, 1804), Dalla Torre (1897), and possibly Schultz (1906) (see above); he apparently accepted Dalla Torre’s synonymies, and his description included characters of both sexes, but even those for males not agreeing with those of the type of ursus (see below). It is thus obvious that Šustera misidentified the type species.

Schulz (1912, mid November) critically reviewed the contributions of the classical Scandinavian authors to knowledge of the Hymenoptera, in particular through examination of all of their type specimens that he could trace. He reviewed all of the works of J.C. Fabricius and quoted the descriptions found in his last relevant work, the Systema Piezatorum (1804), evaluating synonymies where necessary. He noted that Fabricius‘s types were marked by a small, square, green label on the pin, so that they were identifiable as such with some confidence. He noted Nielsen’s (1907) footnote, and also concluded, after repeatedly examining it, that the “type” of Sphex ursus in Copenhagen was a male mutillid and probably a species of Myrmilla Wesmael, 1851. Schulz noted that there were two additional male specimens of ursus in Fabricius’s own collection then in Kiel (but presumably not marked as types), and provided a more detailed description than Fabricius had, in translation: “11 mm long, black and black (nowhere white) hairy. 1^st abdominal segment red, likewise the 2^nd (with the exception of its black end margin); 2^nd abdominal segment with golden-yellow hairs above, black hairs below. The 1^st not petiolate, but triangular or bell-shaped, with a longitudinal keel below. Compound eyes on the inner edge not emarginate.” He was unable to suggest the species’ distribution, except to state that he had not found it in André’s [1899–1902] work on Palaearctic mutillids, nor in Bingham’s [1897] “Fauna of British India”.

Haupt (1927: 271, 308) cited Schulz’s (1912) identification of the type of ursus as a “Mutilla”, but claimed that Šustera’s designation of P. ursus was understood to mean P. rhombica (recognizing Šustera’s misidentification, and probably accepting Dalla Torre’s (1897) suggested ‘synonymy’), and therefore Haupt specified “Paraferreola rhombica Christ” as the type species of Paraferreola. Haupt’s interpretation was almost certainly correct, to judge from the species included in the genus by Šustera (the name rhombica does not appear in Šustera’s paper, nor do the names of any of the other species synonymized with ursus by Dalla Torre); the original descriptions of ursus and rhombica are broadly similar, although differing in a few minor details, so such a misidentification is understandable, and seems to have been generally accepted. Arnold (1935) also gave P. rhombica Christ as the “genotype” of Paraferreola Šustera, and proposed a new monotypic subgenus, Eoferreola Arnold, 1935 with Anoplius soleanus Cameron, 1905 as “genotype”, under Paraferreola.

Pate (1946) referred to Schulz’s (1912) identification of ursus as probably a Myrmilla, and stated that Paraferreola was therefore referable to the Mutillidae; he proposed Tea Pate, 1946 as a “new subgenus” (of Eoferreola) for “Paraferreola Auctt. not of Sustera”, with its type species as “Sphex rhombica Christ, 1791 [= Eoferreola (Tea) rhombica (Christ)]”. Arnold (1948) recounted the situation, and emphasized that “Paraferreola Šustera, being a synonym for a genus of the Mutillidae as yet undetermined, cannot be sustained as a Pompilid genus”, despite the fact that Šustera had misidentified its type species; he considered Eoferreola as a subgenus of Tea based on the fact that Eoferreola had been described as a subgenus of Paraferreola.

Wahis (1986) later designated Tea as a junior subjective synonym of Eoferreola, which meant that Eoferreola became equivalent to Šustera’s original concept of Paraferreola (based on his misidentification). (The confusion in usages of Tea is illustrated in Table 1.) Nevertheless, although Eoferreola has more recently become the preferred name, several recent papers have still used Paraferreola for species of the genus (see Table 2), admittedly mostly in catalogues or lists, but some in non-taxonomic contexts. There are at least four pompilid species (including the type species of Eoferreola) that are still placed in Paraferreola according to the major online aggregator, the Catalogue of Life (Bánki et al. 2021, based on Kroupa and Schmid-Egger 2019, which unfortunately contains many errors according to Christian Schmid-Egger pers. comm.), but the listing of Paraferreola species includes neither rhombica nor ursus. The genus-group names Paraferreola, Eoferreola and Tea have thus all been used for related members of the Pompilidae over more than 100 years, including several recent treatments. However, Pagliano (2008), in a list of Hymenoptera genera and type species, listed Paraferreola (type species Sphex ursus) as an invalid genus equal to Eoferreola (type species Anoplius soleana), and Tea (type species Sphex rhombica) as a junior synonym of Eoferreola.

Despite the fact that various authors (see above) have pointed out that the type specimen of the nominal species Sphex ursus, designated as the type species of Paraferreola by Šustera, is a member of the Mutillidae, the name Paraferreola has never been used as a valid name in Mutillidae. It has either been overlooked, forgotten or perhaps even deliberately ignored, and does not appear in recent surveys of the genus-group names in Mutillidae (Lelej and Brothers 2008; Brothers, Lelej and Williams 2019) nor a checklist of mutillid species (Pagliano et al. 2020), which does not include ursus as a specific name either.

It is evident that confusion about the applicability of Paraferreola in Pompilidae persists, with some authors still using it in that family. In contrast, Paraferreola has never been used as a valid name in Mutillidae. The initial misidentification of the type species is the primary reason for this confusion. Although Haupt (1927) attempted to rectify this, by designating the misidentified species (Sphex rhombica Christ, 1791) which had been accepted as that intended by Šustera as the actual type species, such an action was not in accordance with the rules and practices of zoological nomenclature then applicable, hence Pate’s (1946) proposal of Tea as a new genus-group name with the designation of the same type species. This has not universally been accepted, however, and, additionally, there has been confusion about the relative status of Eoferreola and Tea, although synonymy of Tea with Eoferreola should have simplified matters. However, it is curious that the Catalogue of Life online listing of the species of Eoferreola (Bánki et al. 2021) does not include its type species (Anoplius soleanus), which is instead listed under Paraferreola, so that the species composition of Eoferreola is still questionable; the listing also does not include any mention of Tea as a genus, presumably because it is not considered valid, with several species of Tea all given as valid in Eoferreola. Additionally, Eoferreola rhombica is shown as a senior synonym of Sphex ursus.

The latest edition of the International Code of Zoological Nomenclature (ICZN 1999) provides a solution (Article 70.3.2), which now enables fixation of the species originally intended by Šustera as the type of Paraferreola, something not provided for in previous editions, as follows.

Paraferreola Šustera, 1912. Verh. k. k. Zool.-Bot. Ges. Wien 62: 181 (in key), 200. Male, female.

Type species. Sp[hex] rhombica Christ, 1791 (misidentified as Sphex ursus Fabricius, 1793 by Šustera 1912), by subsequent designation hereby under Article 70.3.2 of ICZN (1999), type locality Europe.

Gender. Feminine.

This accords with the usage intended by Šustera (1912) and accepted by Haupt (1927) and Arnold (1935) in major revisionary works, amongst others. A comment is also needed on the date of publication of Šustera’s work, since it has often been given as 1913. The paper was published in Volume 62 of the Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien, the last (10^th) part of which was only issued in February 1913, so that the volume as a whole was only completed in that year, hence the apparent assumption that the entire content only appeared in 1913. However, the various parts were published separately, and the date given in the volume itself for publication of Heft 5/6, containing pages (129) – (190) and 129–208, is 30^th September 1912, and that for Heft 7, containing pages (191) – (206) and 209–256, is 25^th October 1912. Šustera’s paper spanned pages 171–213, so that all except the last four pages were published on 30^th September 1912. (The dual numbering system apparently differentiated administrative matters, including reports from the various sections of the Academy (page numbers between parentheses) and the scientific papers themselves. The volume is available online at https://www.biodiversitylibrary.org/item/47784#page/5/mode/1up).

The implications of this action are the following: (1) Tea Pate, 1946 is an objective junior synonym of Paraferreola Šustera, 1912; (2) if Wahis’s (1986) subjective synonymy of Tea with Eoferreola Arnold, 1935 is accepted, then Eoferreola also becomes a subjective junior synonym of Paraferreola.

Associated with the continuing confusion about the validity and application of Paraferreola in Pompilidae, is the fact that a pompilid family-group name has been based on Paraferreola, although it has seldom been used, and then essentially only by its original author. Paraferreolini was proposed by Haupt (1927: 262) as a new tribe in the subfamily Homonotinae Haupt, 1927 to include two genera: Paraferreola Šustera, 1912 and Arachnotheutes Haupt, 1927. Later, Haupt (1949) transferred this tribe to the subfamily Pompilinae. Haupt (1950) included in it the subtribes Anopliina, Batozonina and Episyronina, and even later (Haupt 1957) included the subtribe Platyderina. Haupt (1962, published posthumously) placed “Paraferreolini Haupt 1949” (including several genera) in the subfamily Pompilinae, but placed the genus “Paraferreola Šust. 1913” in the subfamily “Platyderinae Haupt 1949” together with the genus Platyderes Guérin-Méneville, 1844; he keyed the females of eight species of Paraferreola, including two new ones. Zonstein and Wahis (2015) referred to Haupt’s (1962) concept of Paraferreolini, but they did not comment on its validity. Later than Haupt’s (1927) proposal of Paraferreolini, Arnold (1937) proposed the tribe Psammoderini for the genera Psammoderes Haupt, 1929 and Paraferreola Šustera, 1912, implying differences of opinion on the relationships of Paraferreola.

Previous ideas on the identity of Sphex ursus Fabricius, 1793 have been biased by an expectation that it is Palaearctic (although no locality information was provided in the original description), hence the suggestion of its being a Myrmilla. A single specimen (Figs 1–2) identified as “Ursus” is still housed in the Natural History Museum of Denmark, Copenhagen, Denmark, the depository of Tønder Lund’s collection, as recorded by Zimsen (1964). Schulz (1912) noted that Fabricius’s type specimens were easily identifiable by having small square green labels on the pins, and presumably found such a label on the Copenhagen specimen he considered to be the type. He also noted that there were two specimens identified as ursus in Kiel (then the repository of Fabricius’s own collection), but those presumably without the green labels. Although the Hymenoptera from Fabricius’s collection have been transferred to Copenhagen on permanent loan, the additional specimens are not there (Lars Vilhelmsen, pers. comm.). The ‘type’ specimen now lacks a small green label but has a typeset red “TYPE” label (probably placed there by Zimsen). It has no locality label (according with the lack of locality information in the original description), but has an identification label (“P. Ursus”), handwritten but reflecting its subsequent placement in Pompilus, and with several pin holes indicating repeated removals and replacements; the handwriting is not that of Fabricius, who generally only gave the specific name and had untidier writing (Lars Vilhelmsen, pers. comm.), but this does not exclude it from being the specimen described by Fabricius, given the considerable amount of time that has passed since its description, and the fact that Fabricius’s original labels had been discarded during reorganization of the collection (Zimsen 1964). Furthermore, Schulz (1912) provided a slightly fuller description than Fabricius had (see above), adding details about the colour and form of the first metasomal tergum and the integumental and setal colours of the second metasomal segment, and those agree fully with the ‘type’ specimen, giving confidence that this is the specimen which Schulz examined. Although Fabricius did not provide any information on the number of specimens he had seen, the fact that he specified Tønder Lund’s collection as the only repository, and provided no locality information, suggests that a single specimen was involved. We thus consider that the specimen in Copenhagen is the holotype (Figs 1–2). It bears the labels as shown in Fig. 3.

Figures 1–3. 

Sphex ursus Fabricius, 1793, holotype, ♂ 1 habitus, lateral view 2 habitus, dorsal view 3 labels.

Examination of photographs of the holotype (Figs 1–2) have shown that it is certainly conspecific with males of Dasymutilla vesta (Cresson, 1865), the most widely distributed species of Dasymutilla Ashmead, 1899 in North America, easily recognized by its smooth protruding subcircular eyes, short entirely punctate tegulae, narrowed campanulate tergum I, and general colour pattern (integument black except for ferruginous tergum II and sternum II, and setation entirely black except pale on tergum II), amongst other characters (Mickel 1928; Manley et al. 2020). The integumental coloration varies somewhat (as shown by figs 281 and 282 in Manley et al. 2020), and the type of ursus has the first tergum also ferruginous and much of the remainder of the integument appearing slightly faded to very dark brown rather than intensely black (perhaps a consequence of its age, though), so being more or less intermediate between the extremes illustrated there.

Although Mutillidae are of biological interest because of their habits and sexual dimorphism, they are of no economic importance, not often seen unless being searched for, and so seldom included in the literature except for specialist taxonomic works or reports of collection records. Although the species involved here is the most widespread species of Dasymutilla in North America, and the name D. vesta is thus applied to many specimens in collections, it is not in very widespread use in the literature, although it meets the conditions specified in Article 23.9.1.2 of the Code (ICZN 1999) for the maintenance of “prevailing usage” (apparently having been used as a valid name in about 30 works published over the last 50 years by at least 22 different authors). However, ursus Fabricius, 1793 has been used as a valid name since 1899, whether for a pompilid (Schulz 1906; Šustera 1912) or a mutillid (Nielsen 1907; Schulz 1912; Haupt 1927; Pate 1946; Arnold 1948) species, and therefore does not meet the requirement of Article 23.9.1.1. which would enable reversal of priority. The major consequence of this identification is that Sphex ursus Fabricius, 1793 must be recognized as a senior synonym of Mutilla vesta Cresson, 1865 and its junior synonyms, as follows, based on Manley et al. (2020), with corrections. (Note that the specific name “ursus”, a bear, is a Latin noun in apposition and therefore unchangeable in gender.)

Dasymutilla ursus (Fabricius, 1793), comb. nov.

Sphex ursus Fabricius, 1793: 210; ♂

Mutilla Vesta (sic) Cresson, 1865: 436; ♀, syn. nov.

Scolia unicincta Provancher, 1882: 6; ♂, syn. nov.

Mutilla monozona Dalla Torre, 1897: 64 (new name for Mutilla unicincta (Provancher, 1882), not Mutilla unicincta Lucas, 1848), syn. nov.

Mutilla sappho Fox, 1899: 239; ♀, syn. nov.

Mutilla agenor Fox, 1899: 245; ♂, syn. nov.

Mutilla zella Rohwer, 1910: 50; ♀, syn. nov.

Pycnomutilla harmoniiformis Rohwer, 1912: 455; ♂, syn. nov.

Dasymutilla errans Rohwer, 1912: 457; ♀, syn. nov.

Dasymutilla bosquensis Rohwer, 1912: 457; ♀, syn. nov.

Dasymutilla ferrugatella Rohwer, 1912: 458; ♀, syn. nov.

Dasymutilla coloradella Rohwer, 1912: 458; ♀, syn. nov.

Dasymutilla coloradella virginica Rohwer, 1912 :459; ♀, syn. nov.

Dasymutilla coloradella kamloopsensis Rohwer, 1912: 459; ♀, syn. nov.

Dasymutilla texensis Rohwer, 1912:460; ♀, syn. nov.

Dasymutilla mesillae Rohwer, 1912:461; ♀, syn. nov.

Dasymutilla carolina Rohwer, 1912:462; ♀, syn. nov.

Dasymutilla columbiana Mickel, 1928:119; ♂, syn. nov.

We are grateful to Lars Vilhelmsen (Natural History Museum of Denmark, Copenhagen) for information on the holotype of Sphex ursus and arranging for photographs of it. The comments of Christian Schmid-Egger (Germany), James Pitts (Utah State University, USA) and Juanita Rodriguez (Australian National Insect Collection) on the manuscript are much appreciated. Funding and facilities were provided to DJB by the University of KwaZulu-Natal.