Eumenes verticalis Say, 1824

Grandinete, Yuri Campanholo, Noll, Fernando Barbosa & Carpenter, James, 2018, Taxonomic Review of Eumenes Latreille, 1802 (Hymenoptera, Vespidae, Eumeninae) from the New World, Zootaxa 4459 (1), pp. 1-52 : 31-34

Eumenes verticalis Say, 1824 ( Fig. 20 )

Female. Color, head: Black, with yellow marks as follows: wide spot on basal half of clypeus; inter-antennal region and a narrow stripe on basal half of scape and on upper half of gena, adjacent to the compound eyes.

Brownish marks: on the two-third apical region of the mandibles; labrum; apex of the clypeus (on the apical projections) and marks on flagellum, mainly the first three and last two flagellomeres. Mesosoma: Black with yellow marks as follows: a broad band on the anterior region on the dorsal face of the pronotum; small spot on upper half of mesepisternum, adjacent to the mesepimeron; tegula (with a brownish translucent mark on the center region of it); apical two-third of metanotum; small spot on the side of the upper half of the posterior face of propodeum and one wide mark on the basal half of it. Marks on the apical region of femora; almost entirely tibiae; first two tarsi yellow-brownish and the last three dark brownish. Metasoma: predominantly black, with yellow marks as follows: two small spots on each side of the middle region of T1; a transverse band on apex of T1; a relatively broad band on each side of T2 and transverse band on apex of T2–T4 and S2–S5 (narrower on those sclerites and almost absent on S5). Wings hyaline, with the costal, medial and submedial cells darkened yellow. Veins and pterostigma dark brownish.

Pubescence: Golden pubescence covering the body. Head: long pubescence on frons, vertex and occipital region. Basal and lateral regions of clypeus with slightly shorter pubescence, while on center and apical regions it is very short. In oblique view, the pubescence on clypeus generally appears to be whitish. Pubescence on scape moderately long (shorter than on frons). Mesosoma: covered with long pubescence (as long as on frons and vertex). Metasoma: T1 with moderately long pubescence (shorter than on mesosoma); basal region of T2 with pubescence as long as on T1 and becoming shorter towards apex. The other sclerites have very short pubescence with some slightly long on apex.

Surface of integument, head: Clypeus with coarse punctation homogenously distributed. Frons and vertex with coarse punctation, denser than on clypeus. Behind the lateral ocelli the punctation becomes sparser. Gena with weak punctation and very sparse distributed. Mesosoma: Coarse and dense (as on frons) punctation, except on the upper half of mesepisternum which are slightly sparser and on anterior half of the lateral face of the propodeum which is reduced. Metasoma: absent on the basal third of T1 and coarse on the two-third of the apical region, sparser than on mesosoma, but becoming denser on apex region. T2 with coarse and dense punctation, denser on apex. On lateral margins of T2 the punctation becomes sparser than on dorsal face. S2 with coarse and sparse punctation mainly on lateral margins. T3–T5 and S3–S5 with weak and very sparse punctation. T6 and S6 with no evident punctation.

Structure, head: Center of clypeus strongly convex; apical region of the clypeus concave, forming two apical and rounded projections with very weak carinae on apex ( Fig. 70 ). Inter-antennal region longitudinally cariniform, slightly wider than the antennal socket; lateral ocelli closer to each other than to the compound eyes; occipital carina very close to the compound eye next to the mandibles and weakly angled on middle region of gena.

Mesosoma: carina on dorsal face of pronotum weakly developed, while on lateral face it is more developed. Pronotal fovea present; mesepimeron slightly elevated regarding to the mesepisternum level; tegula with posterior projection weakly developed, rounded; posterior face of strongly concave on basal half; carina on upper half of external side of fore coxa. Metasoma: T1 not more than 2.5 times longer than wide; moderately swollen in dorsal view ( Fig. 38 ), with the lateral margins on apical half concave and lateral margins on apex divergent. T2, in dorsal view, longer than wide, with the lateral margins weakly concave ( Fig. 54 ). Apex of T2 with lamella which becomes reduced until completely disappear. Basal angle of T2, in lateral view, rising up abruptly.

Male: Clypeus narrower than females; F11 long, narrow basally, with the apex pointed and surpassing the apical margin of F8 ( Fig. 89 ). Dorsal surface without longitudinal carina. Microscopic erect bristles on ventral surface of F11. S7 flattened apically, with some bristles scattered on the apex.

Male genitalia: Aedeagus as in figure 105a, b. Paramere ( Fig. 105c ) with long bristles on middle region of gonostyle; digitus broad, not reaching the middle region of the gonostyle, covered with short bristles ( Fig. 122 ); volsella with long bristles on ventral margin and short scattered bristles; cuspis with moderately long bristles (longer than on digitus); distal lobe truncate, without evident bristles.

Variation: The specimens from Canada to the central west of the USA have the yellow marks less developed than those from the central east. Color variation is as follows: apex of clypeus with spots on each side or almost entirely yellow, with just a longitudinal black band on the apex; marks on mesepisternum, upper half of posterior face of propodeum and T 1 may be reduced or even absent; small spot on each side of scutellum (one specimen presented a spot only in one side of the scutellum); marks on lateral margins of T 1 may be reduced; brownish marks on the flagellum may be absent. The integument may be violaceous mixed with blackish, better observed on laterally on the mesosoma and metasoma. Some specimens have the pubescence of clypeus whitish in oblique view. The posterior face of the propodeum may have the concavity weaker; and the punctation of T 2 may be slightly weaker and sparser.

We examined all the type series of E. coloradensis Cresson, 1875 ( Fig. 21 ); E. cruciferorum Viereck, 1908 (lectotype female and paralectotype male by present designation); E. enigmaticus Viereck, 1908 (lectotype and paralectotype male by present designation); E. stenogaster Isely, 1917 ; and most of the specimens of E. verticalis coloradensis . All of them presented the same features described for E. verticalis , showing only the yellow marks frequently more developed as follows: clypeus often yellow with a black mark that goes from the base to the middle area and sometime on the lateral margins (or only with the black on center region of it); mark on the scape longer; spots on each side of the scutellum, which vary in size (and sometimes absent); the spots on the sides of the posterior face of the propodeum may be wider and often contiguous (in some species totally absent); the band on the lateral margins of T2 broader and sometimes contiguous laterally with the apical band, which it is also clearly broader, as long as the apical band on the T3–T5 and S2–S6. Although the differences relating to the yellow marks, the pubescence, punctation, and external morphology (and male genitalia) were the same as typical verticalis , all included in the range of variation listed. One specimen from Montana showed the punctation of the basal half of T2 sparser, as seen in E. verticalis from Idaho, Wyoming, and Utah.

Bequaert (1944) described E. verticalis var. neoboreus ( Fig. 22 ) and stated that this variety has the same markings as E. verticalis , but with “white to creamy-white instead of yellow” and “the more northern form of the species and is known only from the Dominion of Canada … In the western provinces, it occurs in some localities together with the typical form and there are also a transitional species” (pag. 87). We examined just one male in addition to the type specimens (for which we did not have authorization to extract the genitalia), and we did not find evident differences in pubescence and morphological structure, including the morphology of the male genitalia. One difference that we did find was a male with the punctation of T2 weaker and sparser than that on the type species, a pattern commonly seen in the East coast of the USA (see below), although this species is recorded from Wyoming state (species of E. consobrinus also presented this pattern on the same region). The morphology of the male genitalia was similar to other genitalia examined (see male genitalia section), which supported placing E. verticalis neoboreus as a synonym of E. verticalis with a color variation possibly due the longitudinal differences between populations. We decided to maintain the term neoboreus to facilitate the recognition of the variation, as we proposed for the other variants below. Among all paratypes of E. verticalis neoboreus deposited in the ROM, one specimen (male) was actually E. fraternus .

As we observed for E. consobrinus , the West coast of the USA has populations with different punctation and color than the typical eastern forms. We observed the same in the material examined of E. verticalis tricinctus Isely, 1917 ( Fig. 23 ), including the following yellow marks: wider marks on mesepisternum, including on the basal half; marks on scutellum wider, often contiguous, occupying the anterior half; marks on the lateral margins of the posterior face of the propodeum more developed, marks on the middle area of T1 more developed, often contiguous laterally with an apical band; lateral and apical bands of T2 more developed, occupying almost the entire sclerite, except for black marks on the basal half (and sometimes also laterally) and a transverse band on the middle area (sometimes the transverse band is contiguous with the basal mark). This pattern of marks was observed mostly in females, while the males generally had the same pattern that we described for the typical species and/or coloradensis variants and the genitalia were not different from the variants of E. verticalis . All the specimens of verticalis variety presented the punctation of T2 clearly sparser than the typical form, mainly on the basal half. The types of E. verticalis tricinctus also presented the mesosomal punctation slightly weak, but most specimens matched the typical form.

Type material. The holotype male of Eumenes verticalis var. neoboreus Bequaert, 1944 is excellent condition and bears the labels: ‘8 La Treppe \ VII 42 Qué. \ J. Ouellet’ ‘[Red Label] M.C.Z. \ holotype \ neoboreus \ ♂ ’ ‘ Eumenes \ verticalis \ var. \ neoboreus \ J. Beq’ ‘[Red Label] M.C.Z. \ Holotype \ 33185’. The four males paratypes deposited at ROM are in good condition and bear the labels: ‘Wm. Brodie \ Collection’ ‘ PARATYPE \ Eumenes verticalis \ neoboreus Bequaert \ Det. Bequaert 1944 \ ROMEnt Spec. No. 81631 \ ROM 003001683’; ‘Toronto, Ont. \ 19.7.94 ’ ‘Wm. Brodie \ Collection’ ‘[red square label]’ ‘ PARATYPE \ Eumenes verticalis \ neoboreus Bequaert \ Det. Bequaert 1944 \ ROMEnt Spec. No. 81633 \ ROM 004000144’; ‘Toronto, Ont. \ 23.1.91 ’ ‘Wm. Brodie \ Collection’ ‘[red square label]’ ‘ PARATYPE \ Eumenes verticalis \ neoboreus Bequaert \ Det. Bequaert 1944 \ ROMEnt Spec. No. 81634 \ ROM 004000145’; ‘Toronto, Ont. \ 2.7.94 ’ ‘Wm. Brodie \ Collection’ ‘[red square label]’ ‘ PARATYPE \ Eumenes verticalis \ neoboreus Bequaert \ Det. Bequaert 1944 \ ROMEnt Spec. No. 81632 \ ROM 004000143’ [this specimen is actually E. fraternus ].

The holotype female of Eumenes coloradensis Cresson, 1875 is in good condition but it is lacking the F3–F10 of the right antenna and bears the labels: ‘Col.’ ‘[Red Label] LectoTYPE \ 2728’ ‘ Eumenes \ coloradensis \ Cress.’. The Lectotype female of Eumenes cruciferorum Viereck, 1908 is lacking the fore and mid right legs, and the last two tarsi of the left fore leg. It bears the labels: ‘Oak Creek Canon \ Ariz. 6000 ft. Aug \ F. H. Snow’ ‘1708a’ ‘[Red Label] Eumenes \ cruciferorum \ Type Vier.’. The paralectotype male of Eumenes cruciferorum Viereck, 1908 is in bad condition, lacking: left antenna; left fore and hind legs (except for the coxa); left mid leg from tibia and beyond; right leg; right mid and hind legs from tibia and beyond; metasoma broken but still pinned with the specimen. Bears the labels: ‘Oak Creek Canon \ Ariz. 6000 ft. Aug \ F. H. Snow’ ‘1708b’ ‘[Red Label] Eumenes \ cruciferorum \ Type Vier.’. The lectotype male of Eumenes enigmaticus Viereck, 1908 is in good condition, except for the lacking of the third tarsus and beyond of the hind righ leg. Bears the labels: ‘Oak Creek Canon \ Ariz. 6000 ft. Aug \ F. H. Snow’ ‘1708c’ ‘[Red Label] Eumenes \ enigmaticus \ Paratype Vier.’. The paralectotype male of Eumenes enigmaticus Viereck, 1908 is in excellent condition and bears the same labels as the lectotype, except for one: ‘1708’ instead of ‘1708c’. The holotype of Eumenes stenogaster Isely, 1917 is in excellent condition and bears the labels: ‘Coll. \ Townsend’ ‘White Mts \ 8-1 NM’ ‘Rio Ruidoso \ Abt 6500 ft.’ ‘On fls Monarda \ stricta ’ ‘[Red Label] Type \ No. 21379 \ U.S.N.M.’ ‘ E. \ stenogaster \ Isely’.

The holotype female of Eumenes tricinctus is in good condition and bear the labels: ‘Oreg \ 2511’ ‘Collection \ CFBaker’ ‘[Red Label] Type \ No. 21383 \ U.S.N.M.’ ‘ E. \ tricinctus \ Isely’.

The holotype female of Eumenes cruciferoides is in good condition and bears the labels: ‘Nogares \ (Orizaba) \ 1540 m. 1.IX.75 \ Giordani Soika’ ‘[Red Label] HOLOTYPE \ Eumenes \ cruciferoides Sss. ’.

Figures

FIGURES 13–26. Eumenes species, lateral view. 13) E. cubensis Cresson, 1865 (female); 14) E. flavitinctus Bohart, 1950 (Holotype, female); 15) E. fervens Say, 1824 (Lectotype, female); 16) E. pictus (Smith, 1857) (female); 17) E. sculleni Bohart, 1950 (Holotype, male); 18) E. smithii de Saussure, 1852 (female); 19) E. belfragei Cresson, 1872 (Lectotype, female); 20) E. verticalis Say, 1824 (female); 21) Eumenes verticalis coloradensis Cresson, 1875 (lectotype, female); 22) E. verticalis neoboreus Bequaert, 1944 (Holotype, male); 23) E. verticalis tricinctus Isely, 1917 (Holotype, female); 24) E. filiformis de Saussure, 1855 (famele); 25) E. rufomaculata Fox, 1899 (Lectotype, female); 26) Eumenes versicolor de Saussure, 1852 (Lectotype, female). Scale bar = 1mm.

FIGURES 66–76. Clypeus. 66) Eumenes americanus (female); 67) E. smithii (female); 68) E. crucifera (female); 69) E. crucifera (male); 70) E. verticalis (female); 71) E. fraternus (female); 72) E. fraternus (male); 73) E. sculleni (female); 74) E. filiformis (female); 75) E. rufomaculatus (female); 76) E. versicolor (female). Scale bar = 1mm.

FIGURES 27–42. First tergum, dorsal view. 27) Eumenes americanus; 28) E. bequaerti; 29) E. bollii; 30) E. consobrinus; 31) E. coyotae; 32) E. crucifera; 33) E. cubensis; 34) E. flavitinctus; 35) E. fraternus; 36) E. pictus; 37) E. sculleni; 38) E. smithii; 39) E. verticalis; 40) E. filiformis; 41) E. rufomaculactus; 42) E. versicolor. Scale bar = 1mm.

FIGURES 43–58. Second tergum, dorsal view. 43) Eumenes americanus; 44) E. bequaerti; 45) E. bollii; 46) E. consobrinus; 47) E. coyotae; 48) E. crucifera; 49) E. cubensis; 50) E. flavitinctus; 51) E. fraternus; 52) E. pictus; 53) E. sculleni; 54) E. smithii; 55) E. verticalis; 56) E. filiformis; 57) E. rufomaculactus; 58) E. versicolor. Scale bar = 1mm.

FIGURES 77–92. First tergum, dorsal view. 77) Eumenes americanus; 78) E. bequaerti; 79) E. bollii; 80) E. consobrinus; 81) E. coyotae; 82) E. crucifera; 83) E. cubensis; 84) E. flavitinctus; 85) E. fraternus; 86) E. pictus; 87) E. sculleni; 88) E. smithii; 89) E. verticalis; 90) E. filiformis; 91) E. rufomaculactus; 92) E. versicolor. Scale bar = 1mm.

FIGURES 102–109. Male genitalia: a, b) aedeagus (lateral and ventral view, respectively); c) paramere, internal view. 102) E. pictus; 103) E. sculleni; 104) E. smithii; 105) E. verticalis; 106) E. filiformis; 107) E. rufomaculatus; 108) E. versicolor; 109) General scheme of the apical projection of the aedeagus: a) weakly concave/truncate; b) strongly concave. Scale bar = 1mm.

FIGURES 110–125. Male genitalia: digitus. 110) Eumenes americanus; 111) E. bequaerti; 112) E. bollii; 113) E. consobrinus; 114) E. coyotae; 115) E. crucifera; 116) E. cubensis; 117) E. flavitinctus; 118) E. fraternus; 119) E. pictus; 120) E. sculleni; 121) E. smithii; 122) E. verticalis; 123) E. filiformis; 124) E. rufomaculatus; 125) E. versicolor.