Howard E. Evans.
Studies on Neotropical Pompilidae (Hymenoptera). IX. The Genera of Auplopodini.
Psyche 80:212-226, 1973.

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STUDIES ON NEOTROPICAL POMPILIDAE
(HYMENOPTERA). IX.
THE GENERA OF AUPLOPODINI*
BY HOWARD E. EVANS
Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138, U.S.A.I
The posthun~ous paper of Hermann Haupt on the classification of the Macromerinae (Haupt, 1959) is an unworthy memorial to its author and an unfortunate step backward in the systematics of spider wasps. Working from very little material and a lack of awareness of research in other parts of the world, Haupt erected 12 new genera, few if any of which are likely to stand the test of time. Two of them actually belong in the subfamily Pompilinae, as synonyms of Priochilus Banks (Evans, 1966), while others will fall in the Pepsinae in most classifications (Compsagenia, Anapriocnemis). His inclusion of such diverse elements in the Macromerinae suggests the difficulty in defining the group, which I would rank as a rather weakly characterized tribe of Pepsinae and call the Auplopodini, after the first genus to be used in a suprageneric sense, Pseudagenia, now properly called Auplopus.
I am not in a position to straighten out all the confusion caused by Haupt's paper, but I wish to consider seven genera which he described from the neotropics, all of which can be promptly rele- gated to synonymy. There are, however, several remarkable new genera and subgenera of this tribe in South America, which both Haupt and Banks (1946) were unaware of, and I shall use this opportunity to describe these taxa and to present a revised key to neotropical Auplopodini.
I am much indebted to Professor J. 0. Husing, of the Zoolo- gisches Institut, Halle, for permitting me to borrow some of the specimens that Haupt studied, including several types. Pseudageniella Haupt
Pseudageniella Haupt, 1959, pp. 23, 46 (type-species: P o ~ u s rusticus Fabricius, 1804, monotypic and by designation). *Published with the aid of a grant from the Museum of Comparative -
Zoology.
'Present address: Dept. of Entomology and Zoology, Colorado State Univ., Fort Collins, Colo. 80521.
Manuscript received by the editor, June 7,1973. Psiffte 80:212-226 t 1973). http:ffpsycb.cnlclub.or@~O-212 html



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19731 Evans - Neotropical Pompilidae 213 I have not seen the type of rusticus, but the specimens that Haupt had before him agree perfectly with the type of Priophanes insolens Banks, 1946, and they are from the same locality. This is a well- marked and evidently common species, and these specimens agree well with Fabricius' description. I therefore do not hesitate to con- sider insolens Banks a synonym of rustica Fabricius (new syn- onymy). Townes ( I 957) places insolens in Ageniella, subgenus Ameragenia, an assignment with which I concur; thus Pse~da~eniella Haupt falls as a synonym of Ameragenia Banks, 1945 (new syn-
onymy). In his key, Haupt states that this genus is from the Ne- arctic region, an obvious error, since he states on p. 46 that his material is from Brazil.
Allagenielta Haupt, 1959, pp. 23, 46 (type-species: Allageniella obsoleta Haupt, 1959, monotypic and by designation). Haupt separated this genus from the preceding on exceedingly minor characters, and in fact his lobsoleta and his specimens of rustica differ primarily in size and coloration. I have compared the type specimen of obsoleta Haupt with that of Priophanes plagosa Banks, 1946, and found them to be conspecific; again, both are from the same locality. Townes (1957) also places plagosa in Ageniella, subgenus Ameragenia. Thus Allageniella is to be regarded as a synonym of Ameragenia Banks, obsoleta a synonym of plagosa Banks ( new synonymies).
Brachyagenia H aup t
Brachyagenia Haupt, 1959, pp. 25, 59 (type-species: Brachyagenia nigra Haupt, 1959, monotypic and by designation). By comparison of type specimens, B. nigra is to be regarded as a synonym of Ameragenia thione Banks, 1946;Brachyagenia thus rep- resents still another synonym of Ameragenia Banks (both new syn- onymies ) .
Parageniella H aupt
Parageniella Haupt,1959, pp. 26, 61 (type-species: Priocnemis rufofemoratus Taschenberg, 1869, monotypic and by designation). The type-species is a well-marked Argentinian form and I have little doubt that Haupt and Banks identified it correctly. Banks placed rufofemorata in Priophanes, but like most of Banks' South



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214 Psyche
[September
American Priophanes the species will run to Ameragenia in Townes' ( 1957) key. I would regard Parageniella as still another synonym of Am eragenia (new synonymy).
Cosmagenia Haupt
Cosmagenia Haupt, 1959, pp. 28, 63 (type-species: Agenia amabilis Tas- chenberg, 1869, by designation).
I have studied Haupt's material of amabilis and found it con- specific with Ageniella amoena Banks, 1946. Since Haupt presum-
ably had access to Taschenberg's types, it seems safe to place amoena in the synonymy of amabilis (new synonymy). This species is prop- erly placed in the genus Priocnemella Banks, 1925, and Cosmagenia can thus be relegated to the synonymy of that genus (new syn- onymy)
Compsagenia Haupt
Compsagenia Haupt, 1959, pp. 29, 66 (type-species: Compsagenia laevipes Haupt, 1959, monotypic and by designation). Study of the type specimen of laevipes shows it to be conspecific with the type of Nannochilus ~obscurus Banks, 1946 (new syn- onymy).
Townes ( 1957) has correctly placed Nannochilus in the synonymy of Minagenia Banks, I 934, and Compsagenia Haupt should now be added to the list
(new synonymy). Since levipes
Cresson, 1869, also belongs to this genus, Haupt's laevipes is both a homonym and a synonym. Townes assigns Minagenia to the sub- family Ceropalinae, tribe Minageniini. I would assign it to the Pepsinae, but I do not pretend to understand how the genera should be grouped within that subfamily; at any rate Minagenia does not belong in the Auplopodini.
Anapriocnemis Haupt
Anapriocnemis Haupt, 1959, pp. 25-26, 60-61 (type-species: Pompilus flavipes Guerin, 1836, by designation).
P. flavipes is a well known Chilean species which has been as- signed by Townes (1957) to the genus Priocnemis, subgenus Sphic- tostethus Kohl, along with the other two species assigned by Haupt to Anapriocnemis. Since the type of Sphictostethus has a brachypter- ous female, it might be argued that flavipes is not correctly assigned to that subgenus (though I would not so argue) ; in any case Ana- priocnmis may be laced in the synonymy of Priocnemis Schiodte,



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19731 Evans -- Neotropical Pompilidae 215 which is placed in the Pepsini in Townes' classification (new syn-
onymy).
Mystacagenia, new genus
Type-species. - M. variegata, new species Generic characters. -Females with the general features of Age- niella, including wing venation (shown in Fig. 2) ; males unknown. Length of known species 5.5-8.5 mm; variously colored, with banded wings. Maxillary palpi very long, capable of reaching apex of front coxa; mentum with a few thin setae; mandibles with a basal swell- ing just below which there is a group of long, pale setae which overly and partially conceal the mandible; mandibles slender apically, with a small tooth on the inner margin that is strongly set off from the shaft. Labrum partially exposed, with a deep median notch; clypeus about as wide as lower face; malar space well developed, at least half as long as width of mandibles at their base; antennae unusually slender (Figs. I, 3). Propodeum sloping evenly, its surface smooth and devoid of setae; legs devoid of setae except for very minute ones on the tibiae and tarsi; claws dentate. First abdominal segment hour- glass shaped at extreme base; last segment bearing a number of setae, without a smooth pygidial area.
Key to species (females)
I. Front of head, including mouthparts, white in color; vertex bearing several strong, curved, white setae just behind the ocelli (Fig. 3) ; hind wings with a subapical band ........ .................................................................. albiceps, new species Front o'f head largely ferruginous; vertex without prominent ................................................. setae; hind wings hyaline 2
2. Abdomen largely fuscous except basal and apical segments white; fore wings hyaline, with a strong band below the stigma and a narrow band over the transverse median vein; length of fore ........................................... wing 5 mm
bellula, new species
Abdomen with all segments irregularly blotched with brown and white; fore wings opaque whitish, crossed by three brown bands, the outer two connected above; length of fore wing 7.5 mm .................................................... variegata, new species Mystacagenia variegata, new species
Holotype, - 9, Nova Teutonia, Santa Catarina, Brazil, 2 I Jan. 1956 (Fritz Plaumann) [Coll. H. K. Townes, Ann Arbor, Mich.].



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Psyche [September
Fig. 1. Anterior view of head of Mystacagenia variegata n. sp., type Q. Fig". 2. Wings of same specimen color pattern not shown. Fig. 3. Anterior
view of head of M. albiceps n. sp., type 9. Fig. 4. Wings of Dimorpha- (lema naumanni n. sp., allotype 9. Fig. 5. Mandible of same specimen. Fig 6. Mandible of Agemella (Cyrtagenia) innuba n. sp., type $. Fig. 7. Lateral view of head of same specimen. Fig. 8. Subgenital plate of Pi- morphagenia naumanni n. sp., type
$. Fig. 9. Genitalia of same specimen,
ventral aspect.




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19731 Evans - Neotropical Pompilidae 217 Description. - Length 8.4 mm; fore wing 7.4 mm. Head light rufous, with a pair of black spots on upper front; front and temples with a reticulate pattern of white; malar space, mandibles, and labrum mostly white; thoracic dorsum mainly light rufous, sides with broad streaks of light rufous and white, also some black on posterior parts of mesopleura and metapleura; propodeum white, with a pair of broad, longitudinal black bands; abdomen mainly whitish or some- what cream in color, tergite I with a small amount of black latero- basally and medioapically, tergites 2-5 with much black basally, ter- gite 6 mostly pale; venter mostly pale, but all pale markings of abdomen irregularly tinged with brown; antennae stramineous and partially infuscated on basal 0.3, again at basal vein, this band con- nected through the first submarginal cell with another band partially crossing the wing at the second submarginal; hind wings hyaline. Body with pale, inconspicuous pubescence; erect setae absent except on clypeus, mouthparts, venter and apex of abdomen. Clypeus 3 X as wide as high, its apical margin sinuate, with a broadly rounded median lobe ; front broad, middle interocular dis- tance .60 X head width; upper interocular distance .70 X lower interocular distance; vertex very weakly arched between eye tops, subcarinate behind ocelli; postocellar line: ocello-ocular line = 4:5 ; antennae very slender, third segment 6.6 X as long as its apical width, I .2 X upper interocular distance (Fig. I ) . Pronotum rather flat dorsally, its posterior margin broadly angulate; postnotum widened at the midline; wing venation shown in Fig. 2. Mystacagenia bellula, new species
Htolotype. -9, Avispas, 30 mi. from Marcapata, 'Cusco, Peru, 1-15 Oct. 1962 (Luis Pena) [Coil H. K. Townes, Ann Arbor, Mich.].
Description. - Length 6.2 mm; fore wing 4.7 mm. Head testa-
ceous to somewhat orange, with small dark blotches at center of inner eye margins and a larger dark blotch in front of anterior ocellus; temples, malar space, and area below antenna1 sockets more or less white; mandibles, labrum, and palpi white; thorax and pro- podeum rufous except propleura and anterior corners of pronotum white; abdomen dark brown except all of segments I and 6 and much of sternite 5 contrastingly white; scape white, flagellum white on basal half, testaceous on apical half, streaked with fuscous on lower surface throughout; legs white, with a complex pattern of dark
brown and a small amount of
rufous at base of middle and hind




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218 Psyche [September
coxae.
Wings clear hyaline, fore wing with a brown band over transverse median vein (weakly extended along basal vein) and a much broader brown band nearly crossing wing at stigma. Body pubescence very fine and inconspicuous; body with scarcely any erect hairs except for those on mandibles, a few on clypeus, and some thin ones on apex and venter of abdomen.
Clypeus 2.7 X as wide as high, shaped as in the preceding species; front less broad than in variegata, middle interocular distance .55 X head width; upper interocular distance .73 X lower interocular dis- tance; vertex very weakly arched above eye tops; postocellar line: ocello-ocular line = 6:7; antennae very slender, third segment 6 X as long as its apical width, 1.2 X upper interocular distance. Pro- notal disc rather flat, very short, posterior margin broadly angulate; metanotum angularly projecting backward medially; midline of pro- podeum weakly impressed; wing venation differing from that of variegata in no important details.
Mystacagenia albiceps, new species
Holotypc. -9, Avispas, 30 mi. from Marcapata, Cusco, Peru, 1-15 Oct. 1962 (Luis Pefia) [Coll. H. K. Townes, Ann Arbor, Mich.].
Description. - Length 5.6 mm; fore wing 4.5 mm. Head and mouthparts entirely white except vertex and occiput blotched with testaceous; thorax and propodeum predominantly rufotestaceous, blotched with fuscous across much of pronotal disc, center of meso- scutum, base and apex of scutellum, along pleural sutures, and over most of venter; abdomen rufotestaceous, blotched with fuscous on sides of tergites 1-4, tergites 2 and 3 also with small lateral white spots; scape mostly white, flagellum brown, darkened toward apex; legs mostly rufotestaceous, coxae blotched with fuscous, apices of femora and most of tibiae blotched with fuscous and annulated with white. Wings hyaline except hind wing with a preapical brown band, apex clear; fore wing brownish at base, across basal and transverse median veins, and in a broad band below stigma, the last band ex- tended along radial vein. Pubescence delicate, inconspicuous ; clypeus with several white setae in addition to the tufts on the mandibles, ocellar area also with several strong, curved, white setae; scutellum and apex and venter of abdomen with sparse, weaker setae. Clypeus 2.4 X as wide as high, apical margin weakly convex; head subcircular in anterior view; middle interocular distance .60 X head width; upper interocular distance .68 X lower interocular dis-



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19731 Evans - Neotropical Pompilidae 219 tance; vertex strongly elevated above eye tops, especially at ocellar triangle ; postocellar line : ocello-ocular line = 2 : I ; third antenna! segment 7.5 X as long as its apical width, 1.1 X upper interocular distance (Fig. 3). Pronotum short, disc sloping and with no flat dorsal surface; postnotum transverse, not projecting backward medi- ally. Wing venation similar to that of variegata but stigma unus- ually wid,e, third submarginal cell smaller, only 1.5 X as wide as high, removed from wingtip by twice its own width. Dimorphagenia, new genus
Type-species. - D. naumanniJ new species. Generic characters. -With the general features of the Auplo- podini, including the wing venation (shown in Fig. 4) and the form of the first abdominal segment; length 7-10 mm; wings unhanded, lightly tinged with brown. Female: maxillary palpi of moderate length; mentum with a number of strong setae arising near base and directed forward, much as in Anplopus; mandibles slender, with scattered, strong bristles (Fig. 5) ; labrum wholly concealed ; clypeus not extending under lower margins of eyes; n~alar space about one third as long as width of mandibles at their base; temples well de- veloped, not strongly receding, nearly as wide as eyes; vertex ex- tended well above eye tops; ocellar triangle located well before vertex crest; propodeum with smooth contours, slope low and even; legs relatively smooth, but middle and hind tibiae bearing numerous spines of moderate length; claws dentate, tooth arising rather close to outer ray; apical tergite with a flat pygidial area which is devoid of setae but is minutely punctate and shagreened. Male (Fig. 10) : head remarkably enlarged, much wider than thorax, vertex far above eye tops and ocelli, temples much wider than eyes ; malar space about half as long as width of mandibles at their base; antennae elongate, capable of reaching middle of abdomen; tarsal claws and spines of tibiae as in female; first abdominal segment much expanded from the base, but with no evidence of a lateral seam. Subgenital plate tongue- shaped, midline only weakly elevated (Fig. 8) ; genitalia with the basal hooklets absent, parameres elongate, digiti broad and abruptly truncate apically (Fig. 9).
Remarks. -This genus is most closely related to Auplopus, dif- fering in the less strongly petiolate abdomen (especially in the male), the less well defined pygidial area, presence of a short malar space, broad clypeus with a slightly concave apical margin, and several other features. The male genitalia differ in no important details from those of Auplopus.




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Fig. 10. Dimorphagenia naumanni, type 8, wings omitted. Dimorphagenia naumanni, new species
Holotype. - d, Limoncoche, Prov. Napo, Ecuador (OOO 24'8, 76O 36'W) 7 May 1971 (Martin G. Naumann, nest 2048) [Mus. Comp. Zool., no. 32 1051.
Description 'of male type. - Length 7 mm ; fore wing 6 mm. Entire body testaceous except center of front and (to a lesser degree) vertex blotched with medium brown; legs wholly testaceosus; an- tennae testaceous darkened to medium brown beyond basal third, flagellar segments narrowly ringed with fuscous apically. Wings very lightly tinged with brown; stigma testaceous. Pubescence pale, inconspicuous. Body largely devoid of erect setae except for strong bristles on clypeus and mandibles, scattered setae on front, vertex, and thoracic dorsum (but not propodeurn), and numerous short setae toward apex of abdomen.
'Clypeus 2.8
X as wide as high, apical margin weakly concave; middle interocular distance .68 X head width, 1.4 X eye height; upper and lower interocular distances subequal; postocellar line:



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19731 Evans - Neotropical Pompilidae 22 I ocello-ocular line = 2:5; in lateral view, distance from eye tops to vertex crest .7 X eye height, temples about 1.5 X eye width; third antenna1 segment 4.3 X as long as wide, equal to slightly less than half upper interocular distance. Maximum width of thorax only .7 that of head; pronotum weakly expanded dorsally, its midline de- pressed. Wing venation as in female; terminalia as figured (Figs. 8, 9) ; lateral view of body shown in Fig. 10. Allotype. - 9, same data as type except dated 3 July 1971 [Mus. Comp. Zool.] .
Description of female allotype. - Length 10 mm ; fore wing 8 mm. Head, thorax, and propodeum dark brown, somewhat shin- ing; abdomen rufous except base of first segment black; antennae dark brown; coxae, trochanters, and tarsi dark brown, femora and tibiae rufous. Wings tinged with yellowish brown ; stigma light brown. Pubescence cinereous to light brown, rather conspicuous on coxae, pleura, and propodeum. Body with fairly numerous pale, erect hairs, including some on thoracic dorsum and pleura, pro- podeum, coxae, and especially the abdominal venter; hind femora with scattered short hairs.
Clypeus 2.5 X as wide as high, its apical margin weakly concave; middle interocular distance .63 X 'head width, I .I X eye height; upper interocular distance very slightly exceeding lower interocular distance; vertex broadly rounded off well above eye tops, distance from posterior ocelli to top of vertex exceeding po'stocellar line; postocellar line : ocello-ocular line = 2 :5 ; temples strong, although roundly contracted from behind the eyes, in lateral view not quite as wide as eyes; antennae not especially elongate, third segment only about half the upper interocular distance. Maximum width of thorax only slightly less than that of head; features of pronotum and post- notum as described for male; wing venation as in Fig. 4; legs and abdomen as described under generic heading. Paratypes. - 2 $9, same data as allotype [U.S. Nat. Mus., Brit- ish Mus.].
Variation. - Both paratypes are slightly smaller than the allotype (fore wing 7.3, 7.5 mm) but there are no differences worthy of note. Remarks. - Despite the great difference in head structure in the two sexes, there is close agreement in all other essential features, and there can be no question that these are male and female of one species. This is the only case known to me in the Pompilidae in which sexual dimorphism involves a major difference in head size. In this connection the following notes provided by Martin G.



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222 Psyche [September
Naumann may be of interest (his nest no. 2048 ; see type designation for locality) .
This was a nest of Stelopolybia sp., a social vespid that typically nests in cavities. In this case the nest occupied several cavities inside a large carton ant nest (Azteca sp.) attached to a tree trunk, 2 m above the ground. On May 7, a wasp was seen walking about on the surface of the ant nest. It was captured and proved to be a male pornpilid ( the type of this species). On June 2 I both wasp and ant nest were heavily damaged by children, but on July 3 both wasps and ants were still active, and the nest was harvested by chloro- forming it and catching the contents in a sac. The three female pompilids were found among the vespids, the ants, and the rubble. Structure of the females suggests strongly that they build mud cells: this is the usual function of stiff bristles on the labium and a smooth pygidial plate. In this instance it seems probable that they were utilizing a part of one of the cavities inside the ant nest and being tolerated by the ants and the vespids. I suggest that the large head of the male may enable it to pass as a worker Azteca ant. These ants are polymorphic, and the larger workers commonly are macrocephalic. In this instance the workers were considerably smaller than the male Dimorphagenia, but they were of a similar pale color and the larger workers decidedly macrocephalic. Presumably macrocephaly does not occur in the female sex because it would render them unable to perform their usual hunting and nest-building activities. Macrocephaly in the male suggests that the male is more than a passive inhabitant of the nest; perhaps the presence of several such males inhibits attacks by ants and social wasps. One can only
hope tlhat the relationships of these insects can some day be worked out in detail.
Genus Ageniella Banks
Cyrtagenia, new subgenus
Type-species. - Anzeragenia fallax Arli. Subgeneric characters. - Females with the general features of Ageniella s. str. except as follows (males unknown). Mandibles un- usually broad, with a small tooth located close to the apex (Fig. 6) ; clypeus with rather sharp anterolateral corners and with a median, apical angulation ; front, in lateral view, either abruptly subangulate a short distance above the antennal sockets, then flat to the vertex crest, or flattened all the way from the antennal sockets to the vertex (J?ig. 7), in either case with a median prominence just above the



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19731 Evans -- Neotropical Pompilidae
223
sockets. Pronotum short, with a somewhat flattened dorsal part; pro- podeum with smooth contours, without erect setae or with a very few setae on each side; legs relatively smooth, but middle and hind tibiae with several rows of very small spines; brush on inner side of hind tibia continuous to apex. Third submarginal cell receiving second recurrent vein .4 the distance from the base; anal vein of hind wing reaching media well before cubital fork. Known species
with the wings unhanded, the antennae dark but with a white annu- lus near the middle.
Remarks. - Arnold ( 1934) described a genus from Africa in which the female has the front more or less angulate in profile, Arpactomorpha. However, in this genus the angulate portion has a median groove, and below the angulation there is an oblique im- pression on each side of the face.