Paranysson, Guerin-Meneville, 1844

Pulawski, Wojciech J., 2022, A Revision of the Genus Paranysson Guérin-Méneville, 1844 (Hymenoptera: Crabronidae), Proceedings of the California Academy of Sciences 67 (16), pp. 361-393 : 362-367

Genus Paranysson

Paranysson Guérin-Méneville, 1844:441 (as subgenus of Nysson). Type species: Nysson abdominalis GuérinMéneville, 1844, by monotypy.

Helioryctes F. Smith, 1856:358. Type species: Helioryctes melanopyrus F. Smith, 1856, by monotypy.

Pseudohelioryctes Ashmead, 1899:248. Type species: Pseudohelioryctes foxii Ashmead, 1899, by original designation and monotypy.

RECOGNITION. The following characters place Paranysson within the tribe Miscophini: posterior mandibular margin notched; ocelli round and convex; inner eye margins not emarginate, converging above; antennal socket contiguous with frontoclypeal suture; clypeus not divided by vertical sutures into three parts; anal lobe of hindwing at most as long as half submedial cell; hindfemur thickest near middle, with apex simple.

Within Miscophini, the genus shares the presence of three submarginal cells, the second being petiolate, with Plenoculus, Solierella, and Sphodrotes. It differs from all three in having an emarginate lateral margin of female tergum V (Fig. 1), a previously unnoticed character that differentiates Paranysson from all other Larrinae, and in having the penis valves fused and unusually broadened in apical half, truncate apically (Figs. 43, 50). In addition, Solierella differs from Paranysson by a number of characters (see Bohart and Menke, 1976:294), and the Australian genus Sphodrotes in having the occipital carina effaced before reaching the hypostomal carina, the propodeal enclosure setose (setae at least fine, inconspicuous), sternum I with a posteromedian prominence, and the pygidial plate absent in both sexes. In Paranysson , the occipital carina joins the hypostomal carina just before the latter’s apex, the propodeal enclosure is asetose, sternum I has no apicomedian prominence, and the pygidial plate is present in both females and males. The closest to Paranysson is Plenoculus . In the latter genus, however, the propodeal dorsum is finely granulate or finely ridged and the occipital carina is incomplete below ( Bohart and Menke, 1976:294). In Paranysson the propodeal dorsum is areolate or reticulate, and the occipital carina meets the hypostomal carina (other differences given by these authors separate only some, but not all Paranysson ).

Male genitalia are nearly identical in most species ( Fig. 50 ), except that in P. oscari the apical part of penis valve has a spine-like lateral process ( Fig. 43 ).

According to Lomholdt (1985), Paranysson shares with Plenoculus the gonocoxite (as gonostyle) constricted subapically, and penis valves coalesced and expanded apically. This statement is poorly substantiated: in some Plenoculus , e.g., P. cockerellii W. Fox , and P. palmarum F. Williams , the gonocoxite is not constricted subapically (see Figs. 70-72 in Williams, 1960), and at least in P. propinquus W. Fox the penis valves are not expanded apically and not coalesced (Fig. 82 of Williams, 1960).

TAXONOMIC HISTORY. Guérin-Méneville (1844) described Paranysson from Senegal as a subgenus of Nysson , as reflected in its name, obviously because of the petiolate second submarginal cell shared by both. He included only one species, Nysson abdominalis , which thus became the type species of Paranysson by monotypy. F. Smith (1856) was aware of Guérin-Méneville’s description, but did not recognize his species and described a closely related melanopyrus in the new genus Helioryctes . A big confusion followed. Cresson (1882) and Schrottky (1910) described in Paranysson two North American and two Argentinian species, respectively, which are now known to be Zanysson and Metanysson ( Handlirsch, 1887 reported Cresson’s species in the subgenus Paranysson of Nysson ). Ashmead, (1899) believed that his new species foxii from Ethiopia was a female differing from Helioryctes by the absence of a hindcoxal spine or tubercle and thus deserving a new generic name, Pseudohelioryctes ; the unarmed hindcoxa, however, is due to the fact that the specimen is a male (and a junior synonym of Paranysson abdominalis ). In the same paper Cresson transferred from Nysson to Paranysson three North American species (for a total of five), now classified as Zanysson . Handlirsch (1887), Fox (1894), Kohl (1897), and Ashmead (1899) treated abdominalis as a Nysson (subfamily Nyssoninae , now Bembicinae) because they knew it only through the original description. Kohl (1897) and Ashmead (1899), however, recognized the synonymic names Helioryctes and Pseudohelioryctes , respectively, as members of Larrinae. Bingham (1897) transferred Paranysson abdominalis to Helioryctes , assuming the identity of the two genera, but incorrectly used as valid the younger name Helioryctes . R. Turner synonymized Helioryctes with Paranysson in 1912, and Pseudohelioryctes with Paranysson in 1914. Since then, only Paranysson has been used for the genus in the subsequent literature, and no nyssonine has been described in (or transferred to) it.

Kohl (1897:397) thought that Paranysson was just a Nysson with somewhat more developed hindtibial spines. He also thought (1897:388) that Helioryctes (that he knew only through the original description) closely resembled the Australian Sericophorus , whereas Dalla Torre (1897) placed these two genera (and also Sphodrotes ) in the new subfamily Sericophorinae . Turner (1914) accepted this concept, but renamed the subfamily Paranyssoninae (correctly: Paranyssontinae), wrongly thinking that “as Paranysson is an older name than Sericophorus , it should be used for the subfamily”. Modern researchers ( Leclercq, 1968; Bohart and Menke, 1976) observed the morphological similarity between Sericophorinae and Miscophini and combined them into one tribe. Bohart and Menke (1976) provided a detailed description of the genus.

R. Turner (1914) was also the first author to review the six species of Paranysson described by that time by Guérin-Méneville, 1844, F. Smith, 1856, Bingham, 1897, Ashmead, 1899, Cameron, 1910, and by himself in 1912, describing in 1914 another new species, and generating a key for the identification of females (the poorly described P. foxii was excluded). Arnold (1923), in his monograph of southern African Sphecidae , dealt with Paranysson , but his key is no more than a slight modification of Turner’s; in particular, he did not add any new characters. Leclercq (1968), on the other hand, greatly expanded his key, providing a number of previously unused characters, and also including the males and describing two new species (one of which is a junior synonym). He had, however, not seen three species ( P. abdominalis , P. foxii , P. helioryctoides ), and relied on existing descriptions for their characteristics. Some of his characters (e.g., sculpture of the propodeal dorsum, lateral propodeal carina with or without tooth) are variable and do not guarantee a correct identification.

Overall, the papers by the three authors suffered from the following: 1. P. foxii was known to them only through the insufficient original description, 2. the male of P. helioryctoides remained unknown, and 3. populations of P. abdominalis with an all-black thorax were not recognized. The authors also missed some excellent diagnostic features such as the structure of the clypeus, the impunctate mandibular base in P. oscari , and the shape of flagellomere I in the male of P. abdominalis .

NESTING BEHAVIOR. Arnold (1923:13) first noticed that Paranysson nest in sandy soil and that “ P. quadridentatus has a most powerful odor of bugs”. Subsequently Bequaert (1933) observed the nesting habits of P. melanopyrus at Kasenga, Haut-Katanga Province of the Democratic Republic of the Congo. Each nest he studied consisted of a nearly straight, vertical gallery, 6-8 mm wide, sunk in the bare sand to a depth of about 75 cm. The lower end of the gallery branched into a few more or less horizontal cells, placed at about the same level in various directions. Each cell was 10-12 mm long and 4-5 mm in diameter. At the nest entrance there was a mound of loose sand, about 12 cm long, 9 cm wide, and 3 cm high. A curved tunnel ran through the longitudinal axis of the mound; it was slightly wider than the gallery to which it led, and opened at the narrow end of the mound. The sand was more closely packed together and apparently cemented, either by moisture or by saliva, about the tunnel.

The prey found in the cell consisted of various instar larvae of the pentatomid Natalicola pallidus (Westwood) , usually six or seven per cell.

While collecting Paranysson in various African countries, I also noticed this strong odor. Obviously, the source is the pentatomid prey.

Key to species of Paranysson

1. Wing membrane nearly hyaline ( Fig. 11 ); femora black; ocellocular distance 0.30-0.35 × distance between hindocelli ( Fig. 12 ); length no more than 7.0 mm. Oriental, also recorded from southern tip of Arabian Peninsula......................... P. assimilis Bingham

- Wing membrane conspicuously infumate ( Fig. 3 ) except slightly so in some P. helioryctoides ; mid- and hindfemora in vast majority of specimens ferruginous; ocellocular distance 0.40-1.20 × distance between hindocelli; length in most specimens more than 7.0 mm. Africa south of Sahara................................................................... 2

2. Basal portion of mandible impunctate or with only a few, sparse punctures ( Fig. 39 ); clypeal bevel in most specimens markedly inclined between basal part and lamella ( Figs. 37, 38 ), perpendicularly so in some females. Female: hindcoxal spine robust, with appressed setae ventrally, originating closer to anterior margin than to posterior ( Fig. 40 ). Male: palpi conspicuously setose ( Fig. 41 )................................. P. oscari (R. Turner)

- Basal portion of mandible finely, closely punctate; clypeal bevel not markedly inclined toward basal clypeal part. Female: hindcoxal spine, when present, thin, asetose ventrally, and originating at hindcoxa midlength or closer to posterior margin than to anterior. Male: palpi usual, not conspicuously setose..................................................... 3

3. Length of apical flagellomere 2.6 × its basal width ( Fig. 30 ); propodeum without carina between lateral and posterior surfaces ( Fig. 28 ). Female: clypeal free margin without lateral teeth ( Fig. 26 ); hindcoxa unarmed, without tubercle or spine ( Fig. 29 ). Male: terga III-VII to V-VII sparsely punctate; tergum VI with obtuse lateral carina ( Fig. 31 ); pygidial plate narrow, triangular ( Fig. 31 )........................................... P. inermis Leclercq

- Length of apical flagellomere at most 2.2 × its basal width; propodeum with longitudinal carina between lateral and posterior surfaces, carina projecting as triangular tooth at about two thirds of length or broadened there ( Fig. 6 ), exceptionally not broadened. Female: clypeal free margin with lateral teeth (except teeth absent or ill-defined in P. helioryctoides ); hindcoxa either with spine or acute tubercle ( Figs. 6 , 19 , 24 , 35 , 40 , 47 ). Male: terga III-VII closely punctate; tergum VI without obtuse lateral carina, pygidial plate broad, not triangular..... 4

4. Females.................................................................. 5

- Males.................................................................... 9

5. Clypeal bevel separated by carina from basal part ( Fig. 22 ); hindcoxal venter with acute tubercle but without spine ( Fig. 24 )........................... P. helioryctoides R. Turner

- Clypeus without carina between bevel and basal part; hindcoxal venter with conspicuous spine.................................................................... 6

6. Hindcoxal spine originating next to apical coxal margin ( Fig. 19 ); lateral clypeal teeth protruding beyond free clypeal margin between them ( Fig. 17 )....... P. brevispinosus Arnold

- Hindcoxal spine originating at least half midocellar diameter before apical coxal margin ( Figs. 7 , 35 , 47 ); lateral clypeal teeth not protruding beyond free clypeal margin between them ( Figs. 4 , 33 , 45 )............................................................ 7

7. Dorsal length of flagellomere I 1.4-1.6 × apical width; hindcoxal spine originating at or before hindcoxal midlength ( Fig. 35 ); longitudinal carina between propodeal side and posterior surface in most specimens forming sharp tooth at about two thirds of length.................................................................. P. melanopyrus (F. Smith)

- Dorsal length of flagellomere I 2.0-2.4 × apical width; hindcoxal spine originating next to hindcoxal apex, on its inner side; longitudinal carina between propodeal side and posterior surface slightly broadened at about two thirds of length, but in most specimens not forming sharp tooth................................................................ 8

8. Hindcoxal spine as long as hindcoxal venter or longer ( Fig. 47 ); pygidial plate finely, sparsely punctate throughout ( Fig. 48 ); thorax and propodeum all black......................................................................... P. quadridentatus (Cameron)

- Hindcoxal spine shorter than hindcoxal venter ( Fig. 7 ); pygidial plate rugose except basally ( Fig. 8 ); pronotum, scutellum posteriorly, and postscutellum ferruginous in specimens from Senegal, Mali, and Niger ( Fig. 3 ), in some specimens also propodeal enclosure, but all black in those from Ethiopia, Kenya, and Tanzania......... P. abdominalis (Guérin-Méneville)

9. Tergum II impunctate basomedially or with a few sparse punctures ( Fig. 20 ); hindcoxal venter in large specimens with apical tooth on inner side............ P. brevispinosus Arnold

- Tergum II closely punctate throughout; hindcoxa without apical tooth................ 10

10. Flagellomere I concave ventrally ( Fig. 9 ); ocellocular distance equal to distance between hindocelli or slightly greater; pronotum, scutellum posteriorly, postscutellum, and often part of mesopleuron and propodeal enclosure ferruginous in specimens from Senegal, Mali, and Niger (as in Fig. 3 ), but all black in those from Ethiopia, Kenya, and Tanzania.............................................................. P. abdominalis (Guérin-Méneville)

- Flagellomere I slightly flattened ventrally, not or minimally concave; ocellocular distance smaller than distance between hindocelli; thorax and propodeum all black............ 11

11. Free margin of clypeal lobe obtusely angulate laterally ( Fig. 23 ).......................................................................... P. helioryctoides (R. Turner)

- Free margin of clypeal lobe different.......................................... 12

12. Clypeal lamella emarginate mesally, each side of emargination slightly protruding as obtuse tooth ( Fig. 34 ), distance between teeth apices equal to 1.2-1.8 × midocellar diameter; dorsal length of flagellomere I equal to 1.0-1.2 × that of flagellomere II; hindcoxal venter without transverse, preapical carina, all punctate, not concave apically... P. melanopyrus (F. Smith)

- Clypeal free margin gently arcuate mesally, at most with pair of admedian rudimentary teeth ( Fig. 46 ); dorsal length of flagellomere I equal to 1.2-1.6 × that of flagellomere II; hindcoxal venter with preapical, semicircular carina, concave and unsculptured between carina and apex ( Fig. 49 )........................................... P. quadridentatus (Cameron)

Figures

FIGURES 45-50. Paranysson quadridentatus (Cameron). (45) Female clypeus and mandible; (46) Male clypeus and mandible; (47) Female hindcoxa with spine; (48) Pygidial plate of female; (49) Male hindcoxa (arrow shows semicircular carina); (50) Male genitalia in dorsal view.

FIGURES 37-43. Paranysson oscari R. Turner. (37) Female clypeus and mandible; (38) Male clypeus and mandible; (39) Female clypeus and mandibular base in lateral oblique view; (40) Female hindcoxa with spine; (41) Male palpi; (42) Gastral apex of male in dorsal view (arrow shows lateral carina of tergum VI); (43) Male genitalia in dorsal view (arrow shows lateral process of penis valve).

FIGURES 11-15. Paranysson assimilis (Bingham). (11) Whole body of female in lateral view; (12) Ocellar area of female showing large distance between hindocelli; (13) Female clypeus and mandible; (14) Male clypeus and mandible; (15) Female hindcoxa.

FIGURES 3-9. Paranysson abdominalis (Guérin- Méneville). (3) Whole body of female in oblique lateral view; (4) Female clypeus and mandible; (5) Male clypeus and mandible; (6) Posterior part of female mesopleuron and propodeal side (arrow shows the broadening of the longitu- dinal propodeal carina); (7) Female hindcoxa with spine; (8) Pygidial plate of female; (9) Male flagellomere I.

FIGURES 26-31. Paranysson inermis Leclercq. (26) Clypeus and mandible of holotype female; (27) Ocellar area of holotype female; (28) Propodeal side of holotype female in lateral oblique view showing absence of longitudinal carina; (29) Hindcoxa of holotype female showing absence of spine; (30) Apical flagellomere of male; (31) Gastral apex of male in dorsal view (arrow shows lateral carina of tergum VI).

FIGURES 17-20. Paranysson brevispinosus Arnold. (17) Female clypeus and mandible (arrow shows lateral clypeal tooth); (18) Male clypeus and mandible; (19) Female hindcoxa with spine; (20) Basal terga of male.

FIGURES 22-24. Paranysson helioryctoides (R. Turner). (22) Female clypeus and mandible; (23) Male clypeus and mandible (arrow shows lateral clypeal lobe); (24) Female hindcoxa (arrow shows ventral tubercle).

FIGURES 33-35. Paranysson melanopyrus (F. Smith). (33) Female clypeus and mandible; (34) Male clypeus and mandible; (35) Female hindcoxa with spine. 35