Europiella Reuter, 1909

SCHUH, RANDALL T., 2004, Revision of Europiella Reuter in North America, with the Description of a New Genus (Heteroptera: Miridae: Phylinae), American Museum Novitates 3463, pp. 1-59 : 2-12

Europiella Reuter

Type species: Agalliastes stigmosus Uhler.

Europiella Reuter 1909: 83 (n. gen., descr.); Schuh et al., 1995: 379 (n. syn., disc.).

Poliopterus Wagner, 1949: 53 (n. subgen. of Plagiognathus , descr.). (syn. by Schuh et al., 1995: 379).

DIAGNOSIS: Recognized by the relatively small size, length 2.14–3.68; body form varying from elongate and nearly parallelsided to ovoid. Coloration varying from almost completely pale or weakly greenish to almost entirely black, usually with a few dark markings on femora and at bases of tibial spines. Vestiture of dorsum with reclining simple setae and recumbent, weakly woolly, silvery setae; eyes weakly produced laterally, not closely conforming to anterior margin of pronotum. Some species showing strong sexual dimorphism with males elongate, nearly parallel­sided and females strongly ovoid with hemelytra strongly conforming to shape of body. Pale­colored species similar in appearance, including type of vestiture, to Knightopiella , new genus and Megalopsallus spp. Distinguished from both of those genera by the structure of the male genitalia, and from Megalopsallus by the much smaller pygophore in members of that group.

Male genitalia varying greatly in size, but pygophore always moderately large to large relative to total size of abdomen; right paramere truncate apically, with a more or less well­developed protrusion on either side (figs. 3–5), never lanceolate as in most Phylini ; body of vesica with a distinctive curvature and form, and usually with a proximal portion of one strap terminating near midpoint of vesica (see fig. 1, angulata , fig. 2, E. lattini ); vesica always with two apical processes, these sometimes elongate, flattened, and broadened basally (figs. 3–5), and in the form of blades very similar to that found in Plagiognathus spp. , or much shorter and variously acuminate and/or strongly curving and spinelike (figs. 3–5); secondary gonopore large and heavily sclerotized, placed to the side of the chitinous bands of the vesica rath­ er than in the same plane (figs. 3–5), as in Plagiognathus , and distinctly proximad of attachment of spines.

Often breeding exclusively on Artemisia spp. , occasionally on other Asteraceae , and rarely on Lamiaceae in Palearctic.

REDESCRIPTION: Male: Relatively small, elongate to ovoid, total length 2.14–3.68; body in transverse cross­section more or less cylindrical, not flattened. COLORATION (figs. 1, 2): Coloration varying from almost completely white or pale greenish to nearly all black. Mesoscutum often orange laterally. Femora usually with some contrasting dark spots; tibial spines usually black with black bases, sometimes pale but with bases always at least darkened if not distinctly black. SURFACE AND VESTITURE (figs. 1, 2, 7, 9–11): Dorsum smooth, dull, clothed with reclining simple setae and recumbent, weakly woolly, silvery setae. STRUCTURE (figs. 7, 9–11): Head transverse, frons weakly bulging, clypeus generally not visible from above; eyes relatively small, not sexually dimorphic; antennal segment 2 slender, tapering to slightly greater diameter distally, sexual dimorphism very weak; length of labium variable, from just reaching middle coxae to just surpassing posterior margin of hind coxae; claws elongate, gently curving, parempodia always setiform, pulvilli small. GEN­ ITALIA (figs. 3–5): Genital capsule and genitalia moderately large to large relative to size of abdomen; vesica formed of two chitinous bands, always with two apical process, these sometimes elongate and broadened basally, and in the form of blades, or much shorter and variously acuminate and/or strongly curving and in the form of spines; body of vesica with distinctive curvature and form, with a portion of one strap often terminating near midpoint of vesica (e.g., fig. 1, E. angulata , fig. 2, E. lattini ) and most species with a distinctive, broad, membranous projection extending beyond gonopore straps just distad of gonopore (e.g., fig. 4, E. pilosula , fig. 5, E. stigmosa ); secondary gonopore large and heavily sclerotized, placed laterad of the chitinous bands of the vesica rather than between them, slightly proximad of attachment of spines; phallotheca of relatively uniform structure, typical of Phylini (figs. 4, 5); left paramere boat­shaped, typical of Phylini (fig. 3); right paramere distinctive for the group, truncate apically, with a more or less well­developed protrusion on either side (figs. 3–5), never lanceolate.

Female: Elongate ovoid, usually of more robust appearance than male; total body length 2.14–3.32.

DISTRIBUTION: Like many other genera of Phylinae in North America, but with the notable exception of Plagiognathus Fieber , endemism and species diversity in Europiella are restricted almost entirely to the West. Only Europiella decolor Uhler and E. artemisiae (Becker) occur in eastern North America, both species also occurring in the Palearctic.

HOST ASSOCIATIONS: Unlike most members of the Phylini , Europiella spp. are usually found on their hosts when the plants are not in flower. This is true in North America for Artemisia and Chrysothamnus ­associated species, which are collected during the late spring and early summer, whereas the plants flower in the late summer. Europiella spp. in the Nearctic, with the exception of E. lattini , n. sp., are restricted to members of the Asteraceae , primarily Artemisia spp. , whereas in the Palearctic some species are also found feeding on the Lamiaceae .

DISCUSSION: Because sexual dimorphism is pronounced in some Europiella spp. , the two sexes are not always recognizable as belonging to the same taxon. Accurate identifications of females will require association with males, in most cases. Indeed, as with many phylines, placing females to genus can be difficult. For this reason the following key is designed to be used for male specimens, although it will work for females of some species. Information on host associations is often helpful in placing species to genus. Because some aspects of coloration are quite variable within species, positive identification may require dissection of the male genitalia.

KEY TO MALES OF NORTH AMERICAN SPECIES OF EUROPIELLA

1. All antennal segments entirely pale; bases of tibial spines with at most a small, weak, dark area; coloration of dorsum entirely pale............................ 2

— Antennal segment 1 usually entirely black, much less commonly only black near extreme base or on one face; tibial spines with conspicuous black bases; coloration of dorsum variable................ 3

2. Genitalia as in figure 4 (see discussion of color variation under species account) (British Columbia south to Arizona, east to Kansas)............. pilosula (Uhler)

— Genitalia as in figure 5 (northern Arizona).................. signicornis Knight

3. Dorsum entirely pale; femora pale, sometimes with a few darker blotches..... 4

— Dorsum with at least some dark areas, sometimes dorsum almost entirely dark, occasionally dark areas appearing only as faint infuscations of portions of clavus, corium, or cuneus; coloration of femora variable ................................ 5

4. Long, slender species, total length 3.30–3.61, ratio width pronotum/total length 1:3.65; cuneus elongate, as in figure 2; vesica as in figure 4; extreme southern California .............. morrisoni , new species

— Body form not so elongate and slender; cuneus more broadly triangular; vesica variable............................ 8

5. Large species, total length always at least 3.22; vesica as in figure 3; coloration of dorsum almost entirely dark in some populations (fig. 1); widely distributed in western North America consors (Uhler)

— Smaller species, total length always less than 3.08............................ 6

6. Specimens usually with dorsum entirely white, sometimes with some faint infuscation on pronotum and cuneus...... 7

— Males always with some infuscation on dorsum, especially anterior pronotal lobe and areas between veins on hemelytra, giving a mottled appearance (females more frequently almost entirely pale) (fig. 2); vesica with anterior terminal blade very long and decurved (fig. 5); interior western North America....... stigmosa Reuter

7. Genital capsule large and deep, in lateral view appearing more or less parallel­sided and very broad posteriorly; vesica as in figure 4 (British Columbia south to Mojave Desert, east to Colorado)............................ unipuncta Knight

— Genital capsule more slender, in lateral view tapering to a much less broad posterior margin (see fig. 11B for example conformation); vesica as in figure 3 (Oregon to Wyoming)........... carvalhoi Schuh

TABLE 1 Measurements of Europiella Species

TABLE 1 (Continued)

TABLE 1 (Continued) TABLE 1 (Continued)

8. Relatively large species, total length never less than 3.04.................... 9

— Smaller species, total length never greater than 3.50, usually less than 3.05.... 12

9. Length of antennal segment 2 more than 1one­third greater than width of head (Holarctic)........ artemisiae (Becker)

— Length of antennal segment 2 no more than one­fourth greater than width of head... ............................... 10

10. Genitalia as in figure 4 (southern California)................. pintoi , new species

— Genitalia not as in figure 4 for pintoi .. 11

11. Genitalia as in figure 3 (western North America).......... angulata (Uhler)

— Genitalia as in figure 3 (western North America)........... consors (Uhler)

12. Antennal segments 1 and 2 entirely black; narrow, elongate to very elongate species, width of pronotum never greater than 0.99, usually less than 0.95........ 13

— Antennal segment 1 sometimes only partially black, segment 2 always pale on at least distal two­thirds of segment; broader, more ovoid species, width of pronotum always greater than 0.94........... 14

13. Pygophore (and aedeagus) large, diameter of pygophore as measure from side at juncture with pregenital abdominal segments about equal to width of head; coloration of dorsum appearing somewhat mottled; size variation limited, total length apex 2.81–3.05; genitalia as in figure 4 (coastal mountains of Pacific Northwest)...................... lattini , new species

— Pygophore (and aedeagus) smaller, diameter of pygophore as measured from side at junction with pregenital abdominal segments distinctly less than width of head; coloration of dorsum varying from nearly uniform brown to completely pale/white; size variation great, total length 2.14– 3.50; genitalia as in figure 3 (Holarctic, widely distributed in North America)....................... decolor (Uhler)

14. Dorsum weakly to moderately infuscate, appearing mottled, brownish; genitalia as in figure 5 (interior western North America)................... stigmosa Reuter

— Dorsum strongly infuscate, usually appearing blackish, with some contrasting white areas; genitalia as in figure 5 (Great Basin and California south to northern Baja California)............. umbrina Reuter