Eumenes crucifera Provancher, 1888

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 : 16-20

Eumenes crucifera Provancher, 1888

Eumenes globulosus de Saussure, 1855 , 3: 139 [junior secondary homonym of Uespa globulosa Fourcroy, 1785]. NEW STATUS.

Type data: Holotype female at MHNG (examined). Paratype female at USNM.

Type locality: “L’Amérique du Nord ” (Holotype) . “ Colorado ” (Paratype) .

de Saussure, 1875, 254: 101.

Cresson, 1887: 287 (cat.).

Dalla Torre, 1894, 9: 25 (cat.).

Dalla Torre, 1904, 19: 23 (cat.).

Bequaert, 1928, 101: 1004.

Bequaert, 1938, 33: 62 (key), 63.

Bequaert, 1943, 38 (2): 43.

Eumenes crucifera Provancher, 1888: 421 .

Type data: Holotype female at USNM (n° 1977) (examined). Three paratypes (two males and one female) at USNM (examined).

Type locality: “Los Angeles” (Holotype). Two males and one female from California (Los Angeles Co.).

Dalla Torre, 1894, 9: 22 (cat.).

Dalla Torre, 1904, 19: 22 (cat.).

Bequaert, 1938, 33: 69 (note on type).

MacLachlan, 1980, 53: 618, 620 (key).

Buck et al., 2008, 5: 44 (key), 155.

Eumenes (Alpha) marginilineatus Viereck, 1907 , 33: 381.

Type data: Type male at KUNHM (examined).

Type locality: Este’s Park , Larimer County, Colorado ( USA).

Viereck, 1908, 33: 388 (emarginilineatus [!]).

Bequaert, 1938, 33: 69 (note on type).

Eumenes (Pachymenes) bolliformis Viereck, 1908 , 33: 387. NEW STATUS.

Type data: Lectotype, by present designation, female and Paralectotype at KUNHM.

Type locality: “S. Arizona” (both lectotype and paralectotype).

Eumenes robustus Isely, 1917 , 10: 347 (key), 360. NEW STATUS.

Type data: Holotype female at USNM (n° 21382).

Type locality: “ Beulah , San Miguel Co., N.M.”

Bequaert, 1938, 33: 70 (note on types).

Bequaert, 1944, 71: 81.

Bohart, 1951: 884.

Eumenes sternalis Isely, 1917 , 10: 353.

Type data: Holotype female at USNM (n° 21380) (examined).

Type locality: U.S.A. ( Beaver Canyon , Utah).

Bequaert, 1938, 33: 70 (note on types).

Eumenes xanthogaster Isely, 1917 , 10: 359.

Type data: Holotype male at USNM (n° 21381) (examined).

Type locality: U.S.A. ( Los Angeles , California).

Bequaert, 1938, 33: 70 (note on type).

Eumenes bolliformis ; Isely, 1917, 10: 347 (key), 352.

Bequaert, 1938, 33: 68 (note on type).

Eumenes marginilineatus ; Isely, 1917, 10: 363.

Eumenes coloradensis var. (or subsp.) nearcticus Bequaert, 1944 , 71: 76, 82 [replacement name for E. globulosus de Saussure, 1855 , non Fourcroy, 1785].

Eumenes crucifera stricklandi Bequaert, 1944 NEW STATUS.

Eumenes coloradensis var. (or subsp.) stricklandi Bequaert, 1944 , 71: 76, 84.

Type data: Holotype female at MCZ (n° 33184) (examined).

Type locality: “ Cypress Hills , Alberta ” ( Canada).

Eumenes crucifera stricklandi ; Bohart, 1951: 884 (cat.).

Krombein, 1979: 1507 (cat.).

Eumenes crucifera bolliformis ; Bohart, 1951: 884 (cat.).

Linsley, 1962, 55: 153.

Krombein, 1967: 376.

Giordani Soika, 1978, 29: 28.

Krombein, 1979: 1507 (cat.).

MacLachlan, 1980, 53: 618, 621 (key).

Ruiz C. et al., 1993, 88: 86 (list).

Rodríguez-Palafox, 1996: 479 (list).

Eumenes crucifera crucifera ; Bohart, 1951: 884 (cat.).

Krombein, 1979: 1507 (cat.).

MacLachlan, 1980, 53: 618.

Eumenes crucifera nearcticus ; Bohart, 1951: 884 (cat.).

Krombein, 1958: 163.

Krombein, 1979: 1507 (cat.).

MacLachlan, 1980, 53: 618.

Distribution: Mexico (Chihuahua); Canada (Alberta, British Columbia *; Manitoba *, New Brunswick *, New Scotia*, Northwest Territory*, Quebec *, Ontario); USA (AZ, CA, CO, CT*, ID, IL*, IN, KY*, MA*, ME, MI, MN, MO, ND, NH, NJ, NM, NV, NY, OH*, OR*, PE, SC, SD, UT, VT*, WA*, WI, WY).

* Records from the literature.

Comments and diagnosis: Eumenes crucifera Provancher, 1888 ( Fig. 8 ) can be identified by having long and golden pubescence along the body; punctation on the clypeus; black integument with yellow marks varying in size (see variation section below); T2 wide, with the lateral margins strongly concave ( Fig. 48 ); and mainly by the posterior face of the propodeum weakly concave. The species has a wide distribution across the USA and Canada, and consequently we observed a range of variation mainly in coloration, which previous authors used to justify the description of various subspecies.

We had access to all the type specimens of the typical E. crucifera and its subspecies, except for the holotype of E. crucifera flavitinctus , of which we examined four paratypes (females and males). It is important to highlight that one specimen deposited at USNM (n° 21382), labeled as paratype of E. crucifera , is actually a non-type specimen, as Provancher did not mention a paratype from Oregon.

A total of 361 specimens were examined, females and males, and the genitalia from all the subspecies were extracted and examined. As a result, we observed that the same range of variation that occurs in E. verticalis also exists for E. crucifera , which has an extensive geographical distribution that causes intra-specific variation among the specimens, including some overlapping morphological characters. All the variation examined was found in the bolliformis , nearcticus and stricklandi subspecies (the particulars are discussed in the variation section) in various combinations; the male genitalia were compared in each form and presented no significant differences. Therefore, to avoid treating those names as a subspecific category based just on color differences, which is known to be unreliable ( MacLean et al., 1978; Carpenter, 1988; Carpenter and van der Vecht, 1991; Carpenter and Garcete- Barrett, 2003), we propose the synonymy of E. c. bolliformis , E. c. nearcticus and E. c. stricklandi under E. crucifera and treat these names as variations of the typical species as a consequence of the great geographical distribution of this species. Finally, we elevated the subspecies E. c. flavitinctus to specific level based on features that are discussed in the redescription of that species.

Redescription. Holotype Eumenes crucifera Provancher, 1888 ( Fig. 8 )

Female. Color, head: Black, with apex the tooth of mandibles brown. Yellow marks as follows: apex of labrum; clypeus, except for a blackish spot on the center region; inter-antennal region; short and narrow stripe on upper half of the gena, adjacent to the compound eye; broad longitudinal stripe on scape. Mesosoma: Black with yellow marks as follows: anterior portion of the dorsal face of pronotum; wide spot on upper half of the mesepisternum, adjacent to the mesepimeron; tegula (with a translucent spot on center of it); a triangular shaped spot on each side of mesoscutum and a mark on the center region of it; metanotum, except for a narrow stripe on the basal edge; wide spot on each side of upper half of the propodeum; a small spot on mid coxa; distal half of femur, with some brownish marks; entirely tibiae and tarsi, which becomes darker towards apex. Metasoma: T1 black with two big yellow spots on center region and a transverse stripe on apex of T1 and S1. T2 yellow with a black “T” on dorsal face of it. T3–T6 entirely yellow with a transverse narrow brownish stripe on apex. S2 with basal half blackish and narrow stripes on the lateral margins almost reaching the apex; S3–S6 with black marks on basal half. Wings: Hyaline with costal, medial and submedial yellow. Pterostigma and veins brownish.

Pubescence, head: Frons, vertex, gena, scape and occipital region with long and golden pubescence (longer on frons and occipital region). The pubescence on scape is the shortest regarding those other regions. Clypeus with long and slightly whitish (better observed in oblique view) pubescence mostly on basal and lateral regions, while on apex it is shorter. Mesosoma: All covered with golden and long (as on frons) pubescence. Anterior surface of coxae with long and golden pubescence, longer on fore coxa and shorter on hind coxa. Scattered erect golden bristles on trochanter and basal half of the fore femur. Metasoma: T1 covered with golden and long pubescence, longer on basal half. T2 covered with golden pubescence, longer on basal third, becoming shorter towards apex. T3–T6 and S2–S6 covered with short and golden pubescence, except for some long bristles on apex.

Surface of integument, head: Clypeus with and sparse punctation mainly on basal half. Frons and vertex with coarse punctation, moderately dense, becoming weaker and sparser behind the ocelli. Mesosoma: Dorsal face of pronotum, mesoscutum, scutelum and metanotum with coarse and moderately dense punctation, as in frons. Mesepisternum with coarse punctation as mesosucutum but slightly sparser. Posterior face of propodeum with coarse and denser punctation, forming carinae between the punctures. Posterior half from the lateral face of propodeum with coarse but sparser punctation, while on the anterior half the punctation is reduced. Metasoma: Dorsal and ventral face of T1 with coarse and moderately dense (sparser than mesoscutum) punctation mainly on apical two-third; dorsal face of T2 with weaker and sparser punctation regardless T1, and the punctation becomes and sparser towards the lateral face. Basal half of S2 with (more than in dorsal face of T2) scattered punctation. T3–T6 with punctation as in S2, and S3–S6 with no evident punctation.

Structure, head: Clypeus weakly convex, slightly longer than wide with the apex concave, forming two rounded projections with a weak carina on apex of each projection ( Fig. 68 ). Inter-antennal region longitudinally cariniform, as wide as the diameter of antennal socket. Lateral ocelli closer to the compound eyes more than to each other. Occipital carina well developed along all its extension, being slightly angled on middle region of gena.

Mesosoma: Pronotal carina present, weakly developed on dorsal face and stronger on lateral face; pronotal fovea present; mesepimeron slightly elevated when compared to the mesepisternum level; tegula with posterior emargination weakly developed, rounded; parategula lamelliform. Posterior face of propodeum moderately concave on basal half. Fore coxa with a longitudinal carina; posterior face of the propodeum concave on lower half.

Metasoma: T1 about two times longer than wide, strongly swollen in dorsal view ( Fig. 32 ), with the lateral margins subparallel on the distal half, except on the apex which is slightly divergent. Base of T2, in lateral view, rising up abruptly; apical third of T2, in lateral view, with a weak sinuosity before the apex; T2 as long as wide in dorsal view; in dorsal view ( Fig. 48 ), the lateral margins of T2 are strongly concave on middle region; lamella on apex of T2 present.

Male: Clypeus narrower than females ( Fig. 69 ). Sometimes the males of E. crucifera may confused with those from E. verticalis , but they can be rightly separated from them by the absence of the microscopic bristles on the ventral surface of F11. Furthermore, males of E. crucifera present the F11 long, broad basally, with the apex pointed, surpassing the apical edge of F8, sometimes reaching the basal edge of it (in E. verticalis it reaches only the apical edge) (fig. 82). Dorsal surface with a longitudinal carina. Ventral surface without microscopic erect bristles, different from E. verticalis , which have them. The lateral margins of T2 usually are not so concave as on females. S7 flattened apically, with some moderately long bristles scattered on apex (also on T7).

Male genitalia: Aedeagus as in figure 98a, b. The dorsal margim may vary between less to strongly angled, as we observed in E. consobrinus and E. verticalis . Paramere (fig. 98c) with long bristles on middle region of gonostyle; digitus long, surpassing the middle region of the gonostyle, with short bristles on apex, becoming moderately longer towards base ( Fig. 115 ). Distal lobe truncate, without evident bristles. Cuspis with long (longer than on digitus) and erect bristles. Volsella with long bristles (longer than on cuspis) on the ventral edge and scattered short bristles.

Variation, color: The color variation listed was observed also in specimens from the same localities. Clypeus entirely black, except for small marks on each side on the basal half and sometimes on apical half (NM, NV, UT), or entirely yellow; yellow marks on scape, pronotum, mesepisternum, scutellum, T 1 may be reduced or sometimes absent; yellow mark on central area of scutellum may be absent; black mark on T 2 may be covering all the basal half, with just a broad yellow band on each lateral margin; or black mark on S 2 may be covering almost the entire sclerite. Pubescence: Pubescence on apical half of T 2 may be slightly longer (as long as on basal half). Surface of integument: punctation on clypeus may be more evident (specimens from the same localities varied between less and more evident punctures); punctation of T 2 may be stronger, although some specimens outside of California presented punctures as in the typical pattern (NV, UT, ID, WY). Structure: posterior face of propodeum may vary from weakly to strongly concave; generally concave in males. T 1 may be shorter and more developed on apex or longer; lateral margins of T1 more concave; some specimens presented an elevation on the middle region of T2 (better observed in lateral view), which produces a weak sinuosity on the apical third (not so evident as on E. consobrinus ). We observed in two specimens that the sinuosity was strong, but this was clearly caused by a malformation of the sclerite that produced the impression of a strong sinuosity; the lateral margins of T 2 may be less concave, mainly in males; and some specimens may present the basal angle of T2 rising up weakly, as in the type specimen of E. sternalis .

Viereck (1908) described E. bolliformis (lectotype and paralectotype designated herein) ( Fig. 9 ), which was considered by Bohart (1951) as a subspecies of E. crucifera , identified by its ferrugineous color adjacent to the yellow marks along the body (mainly on the mesosoma, T1, and T2), and the coarse punctation of T2.

The type of E. crucifera is from California and shows a pattern of strongly developed yellow marks and the punctation of T2 often weaker. As the distribution goes to the central-east regions, the punctation becomes stronger and the yellow marks often are less developed, although this variation was not a pattern for some specimens from those regions (see variation section). Besides the ferrugineous marks, we also observed for E. bolliformis that the specimens often present the lateral margins of T1 more concave and the posterior face of propodeum also with the concavity more developed. However, this pattern could not justify a subspecies rank because we observed typical crucifera mainly from Utah and Colorado that also showed these features, including some ferrugineous marks adjacent to the yellow marks, apparently like a “transitional species”. As southern specimens were examined, the ferrugineous marks became more developed and the concavity of the posterior face of the propodeum stronger, although it was variable. One specimen identified as bolliformis was recorded from California and as expected it showed more developed yellow marks (with ferrugineous), but the punctation of T2 is weaker. Finally, we compared the genitalia from males of typical crucifera and bolliformis and we did not find any difference that would be enough to separate the subspecies into different species. Therefore, we propose to treat E. bolliformis as a synonymous variant of E. crucifera .

We examined the holotype of E. globulosus ( Fig. 10 ) and, as we observed in E. verticalis , the specimens from Canada and northern to central-west regions of USA presented a pattern with reduced yellow marks on the body. All the variation, especially on T1, the posterior face of propodeum, and punctation of T2, was observed on the specimens identified as E. crucifera nearcticus . As we did not find any pattern to treat it as a different species from the typical species, we propose to treat nearcticus as a variant of E. crucifera . Furthermore, the differences in yellow marks are more visible in females, while males often have the yellow marks reduced and less variable. Some females presented the clypeus with coarse punctation, which may lead them to be confused with E. verticalis (mostly the neoboreus variation), but the latter species still presents the clypeus with punctures clearly more evident, T1 is usually less swollen in dorsal view and the lateral margins of T2 are less concave.

Eumenes crucifera stricklandi ( Fig. 11 ) is known only from the type specimen and besides the yellow-creamy marks on metasoma, there is no difference in pubescence, punctation, and structure compared to typical E. crucifera . We did not examine any males of the subspecies, but propose to treat it also as a variant of E. crucifera .

Type material: The holotype of Eumenes crucifera (female) is in good condition and bears the labels: ‘Los Angeles \ Co. CAL. (C)’ ‘178’ ‘[red label] Type No \ 1977 \ USNM’ ‘[red label] Type \ No. 1977 \ U.S.N.M. [partially handwritten label]’ ‘ E. c. \ crucifera \ Prov.’ ‘DO NOT REMOVE \ SI DB Reference \ Not a property tag \ T. Schultz. NMHN \\ USNM ENT \ 0 0 537193 [bar code]’ ‘Loan from \ USNMNH \ 2073492’. The paratypes at USNM are in good condition and bear the labels: ‘Los Angeles \ Co. CAL. (C)’ ‘[Red Label] Paratype \ No. 21381 \ U.S.N.M.’ ‘ Eumenes crucifera crucifera \ R. Bohart`44 Prov’; ‘Los Angeles \ Co. CAL. (C)’ ‘[Red Label] Paratype \ No. 21381 \ U.S.N.M.’; ‘178’ ‘Los Angeles \ Co., Cal.’ ‘Collection \ Coquillett’ ‘[Red Label] Type No \ 1977 \ USNM’ ‘ Eumenes \ crucifera \ Prov.’.

The holotype of Eumenes xanthogaster (male) is in bad condition with the metasoma broken (glued on a label), and lacking the left F2–F11.

The holotype female of Eumenes globulosus is in bad condition, missing the apical tarsi of the right fore leg, entirely mid right leg, all tarsi of the right hind leg and the flagellum of the both antenna. Bears the labels: ‘Amer. Sept.’ ‘[Red Label] Holo- \ TYPE \ ♀ globulosus \ Saussure’ ‘ MHNG \ ENTO \ 00008850’. The paratype female ( USNM) is in good condition and bears the labels: ‘Ft. Collins \ Col. 1047’ ‘ Eumenes ♀ \ globulosus Sauss. ’ ‘[Red Label] Paratype \ No. 21382 \ U.S. N.M.’.

The holotype female of Eumenes coloradensis stricklandi is in good condition and bears the labels: ‘Cypress Hills, Alta. \ 10 VIII 1939 \ E. H. Stricklandi’ ‘[Red Label] M.C.Z. \ Holotype \ stricklandi \ ♀ ’ ‘ Eumenes \ coloradensis \ var. \ stricklandi \ J. Beq’ ‘[Red Label] M.C.Z. \ Holotype \ 33184’ ‘MCZ-ENT \ 00033184’.

Figures

FIGURES 1–12. Eumenes species, lateral view. 1) Eumenes americanus de Saussure (female); 2) E. belfragei aureus Isely, 1917 (Holotype, female); 3) E. bollii Cresson, 1872 (Lectotype, female); 4) E. bollii oregonensis Bequaert, 1938 (Holotype, female); 5); E. bequaerti Isely, 1917 (Holotype, male); 6) E. iturbide (= E. consobrinus; Holotype, female); 7) E. coyotae Bohart, 1948 (Holotype, male); 8) E. crucifera Provancher, 1888 (Holotype, female). 9) E. bolliformis Viereck, 1908 (Lectotype, female); 10) E. crucifera var. nearcticus Bequaert, 1944 (female); 11) E. crucifera stricklandi Bequaert, 1944 (Holotype, female); 12) E. cruciferoides Giordani Soika, 1978 (Holotype, female). 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 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 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.