Megalopsallus brittoni Knight, 1927: 227

SCHUH, RANDALL T., 2000, Revision of the North American Plant Bug Genus Megalopsallus Knight, with the Description of Eight New Species from the West (Heteroptera: Miridae: Phylinae), American Museum Novitates 3305, pp. 1-70 : 11-21

Megalopsallus brittoni Knight, 1927: 227 (n. sp.) Megalopsallus adustus Knight, 1927: 227 (n. sp.).

NEW SYNONYMY.

DIAGNOSIS: Most similar to femoralis in

reddish brown coloration and spots at bases of setae on dorsum in most specimens, but femoralis more strongly sexually dimorphic. Similar in appearance to nuperus in near absence of sexual dimorphism and protuberant eyes, but that species lacking brown spots at the bases of setae on dorsum. Further distinguished from nuperus by differences in structure of male genitalia (compare figs. 13 and 15).

REDESCRIPTION: Male: Medium sized, total length 3.57–3.82, length apex clypeus–cuneal fracture 2.58–2.71, width across pronotum 1.18–1.29. COLORATION: General coloration, including legs, reddish brown, with some more strongly reddish spots or markings, particularly on head and anterior half of pronotum; eyes distinctly reddish; bases of setae on dorsum often with darkened spots at bases; tibial spines pale with brown bases not strongly contrasting with remainder of tibia. SURFACE AND VESTITURE: Entire body surface dull, matte; dorsum with dark simple setae intermixed with woolly, silvery setae. STRUCTURE: Relatively broad, hemelytra not conspicuously elongate; eyes protuberant; labium long, reaching posterior margin of hind trochanters; tarsal claws elongate, slender, curving, pulvilli very small. MALE GENITALIA: Vesica S­shaped; gonopore subapical; gonopore sclerite not developed (fig. 13).

Female: Total length 3.85–4.16, length apex clypeus–cuneal fracture 2.86–3.10, width across pronotum 1.33–1.42; sexual dimorphism weak, females slightly more robust than males (fig. 7).

HOST: Unknown.

DISTRIBUTION: Known from Connecticut and Texas, primarily from coastal localities.

DISCUSSION: Knight (1927) described brittoni and adustus on the same page, separating them primarily on the length of the second antennal segment. Although this measurement was frequently used by Knight to discriminate species, I have found it to be less reliable than was believed by Knight, owing to variability within species. I have seen only two specimens of brittoni , and have compared them with large numbers of specimens from Texas, including the holotype of adustus from Anahuac, that agree closely with Knight’s concept of adustus . I am treating adustus as a junior synonym of brittoni because both nominal taxa have brown spots at the bases of many of the setae on the dorsum, an attribute found elsewhere in Megalopsallus only in punctatus , the last species otherwise showing little similarity to brittoni .

SPECIMENS EXAMINED: USA. — Connecticut: Stratford, July 9, 1920, B. H. Walden, 1♀ ( USNM). Westville, July 4, 1904, W. E. Britton, holotype: male of brittoni (USNM) . Texas: Brazos Co.: Bryan, April 8, 1965, J. C. Schaffner, 1♀ (TAMU). Cameron Co. : 4 mi ESE of Brownsville, October 20, 1997, W. F. Chamberlain, 13, 1♀ (TAMU). Brownsville , Boca Chica , December 27, 1945, R . H. Beamer, 33, 2♀ ( KU). Chambers Co.: Anahuac, October 8, 1918, H. S. Barber, 3♀ ( USNM); holotype: male of adustus (USNM) . Galveston Co.: Galveston , February 2, 1932, L. D. Tuthill, 23, 1♀ ( KU). Nueces Co. : Corpus Christi , January 1, 1946, R . H. Beamer, 143, 35♀ ( KU, AMNH) .

Megalopsallus californicus , new species

Figures 7 , 13

HOLOTYPE: Male, Schafter, Calif., IX­24­ 1940, G. L. Smith Coll. Deposited in the California Academy of Sciences , San Francisco.

DIAGNOSIS: Recognized by pale green coloration and marmorate membrane, the latter characteristic occurring elsewhere in Megalopsallus only in the heavily reddish brown M. marmoratus Knight.

DESCRIPTION: Male: Relatively small, total length 2.78–3.10, length apex clypeus–cuneal fracture 1.88–2.12, width across pronotum 0.94–1.03. COLORATION: Pale, weakly greenish; eyes red; membrane of hemelytra generally pale, marmorate posterior to cells (fig. 7); femora with some brown spots; tibial spines pale, with at most medium brown bases. SURFACE AND VESTITU­ RE: Dorsum with brown, recumbent, simple setae intermixed with woolly, silvery setae. STRUCTURE: Elongate ovoid; labium long, surpassing posterior margin of hind trochanters; claws elongate, curving, pulvilli small. MALE GENITALIA: Vesica S­shaped, attenuated apically; gonopore relatively small, subapical; gonopore sclerite not developed (fig. 13).

Female: Total length 2.72–2.90, length apex clypeus–cuneal fracture 1.98–2.26, width across pronotum 1.01–1.13; more broadly ovoid than male (fig. 7).

ETYMOLOGY: Named for its occurrence in California.

HOST: Unknown.

DISTRIBUTION: Southern San Joaquin Valley , California .

PARATYPES: USA. — California: Kern Co.: Schafter , September 24, 1940, G. L. Smith, 233, 18♀ ( UCB, AMNH) . Schafter , September 15, 1942, G. L. Smith, 23, 2♀ ( USNM) .

Megalopsallus ephedrae (Knight) , new combination Figures 8 , 14

Merinocapsus ephedrae Knight, 1968: 34 (n. sp.); Schuh, 1986: 218 (diag., distr., figs.).

DIAGNOSIS: Recognized by dull, orange or nearly black head, pronotum, and scutellum and pale to red hemelytra with orange cuneus and smoky membrane; femora always unicolorous orange or dark. Coloration of dorsum often similar to froeschneri , but femora in that species always paler distally than proximally, whereas in ephedrae femora unicolorous over entire length. Head, pronotum, and scutellum in froeschneri usually more strongly polished and shining than in ephedrae .

REDESCRIPTION: Male: Medium sized, total length 3.41–4.32, length apex clypeus– cuneal fracture 2.24–2.81, width across pronotum 0.95–1.14. COLORATION: Head, pronotum, and scutellum dull, usually castaneous to nearly black, sometimes orange; hemelytra ranging from pale to intensely red orange or red, cuneus always orange to red, membrane smoky, veins orange (fig. 8); eyes dark, often mostly black; underside of body usually dark, castaneous; antennae dark; legs of similar coloration to underside of body, tibiae sometimes lighter and tibial spines with visible small dark bases; appendages and underside of body orange in specimens with completely orange dorsum. SURFACE AND VESTITURE: Head, pronotum, and scutellum smooth, dull, clothed with recum­ bent, dark, simple setae intermixed with woolly, silvery setae. STRUCTURE: Elongate, parallel­sided; specimens in some populations relatively longer than those in others; head strongly declivent (fig. 3); labium reaching to about apex of middle coxae; claws moderately elongate, smoothly curving; pulvilli small. MALE GENITALIA: Vesica elongate, twisted, with bifid apex, secondary gonopore subapical with a weakly sclerotized gonopore sclerite (fig. 14).

Female: Total length 2.98–3.22, length apex clypeus–cuneal fracture 2.16–2.34, width across pronotum 1.04–1.12; similar in coloration to males, body not so elongate and more strongly ovoid (fig. 9).

HOSTS: Ephedra apsera , E. nevadensis , E. viridis (Ephedraceae) .

DISTRIBUTION: Northern Baja California , and Mojave Desert of southern California , southern Nevada, and southern Utah.

DISCUSSION: This is the most variable of the Ephedra ­feeding species. The bifid apex of the vesica suggests a close relationship with the Ephedra ­feeding species froeschneri and pallipes .

SPECIMENS EXAMINED: MEXICO. — Baja California Norte: 6 mi E of Ojos Negros, June 9, 1980, Brown and Faulkner, 1♀ (SDNH). 12 mi E of El Rosario, March 25, 1979, J. D. Pinto, Ephedra sp. (Ephedraceae) , 73, 5♀ (AMNH, UCR). USA. — California: Inyo Co. : 2 mi E of Westgard Pass summit, White Mountains, 2125 m, July 2, 1980, R. T. Schuh, Ephedra nevadensis (Ephedraceae) , 6♀ (AMNH). Inyo Mts , May 25, 1937, D. Little, 1♀ (LACM). Rt 95 at North edge of Mono Lake, 2188 m, July 11, 1980, R. T. Schuh and G. M. Stonedahl, Ephedra sp. (Ephedraceae) , 33, 7♀ (AMNH). Tuttle Creek, 2 mi SW of Lone Pine, May 9, 1969, P.A. Opler, 13 (UCB). Mono Co.: 2 airline mi S of Inyo , 8500 ft, June 27, 1961, J. Powell, Ephedra sp. (Ephedraceae) , 43, 2♀ (UCB). Riverside Co.: ca. 0.8 mi N of jct. Deep Creek and Horsethief Creek, T7S R6E Sec 6, 2960 ft, J. D. Pinto, 13 (UCR). San Bernardino Co.: Victorville, April 21, 1935, C.E. Norland, 2♀ (LACM). Providence Mountains State Recreation Area, 4300 ft, May 18, 1982, M. D. Schwartz, Ephedra aspera (Ephedraceae) , 23, 6♀ (AMNH). 10 mi W of Lucerne Val­ ley, 905 m, May 13, 1978, J. D. Pinto and R. T. Schuh, Ephedra sp. (Ephedraceae) , 63, 28♀ (AMNH). 23 mi S of Amboy, April 7, 1966, C. W. O’Brien, 5♀ (UCB). San Diego Co.: Anza­Borrego Desert State Park, Grapevine Canyon, mp 74 on county Rt 52, April 22, 1980, Russell and Schwartz, 13 (AMNH). Nevada: Lincoln Co.: 5 mi NE of jct. of Rts 38 and 93, 2500 ft, May 19, 1982, M. D. Schwartz, Ephedra nevadensis (Ephedraceae) , 263, 18♀ (AMNH). Nye Co.: Northumberland Canyon Rd, Toquima Mts, 15 mi E of Rt 376, T31N R45E Secs 3, 4, 10, 7000 ft, June 29, 1983, R. T. Schuh and M. D. Schwartz, Ephedra sp. (Ephedraceae) , 173, 27♀ (AMNH). Mercury, 17 M, June 12, 1965, H. Knight and J. Merino, Ephedra nevadensis (Ephedraceae) , Paratypes: 33, 6♀ (CNC, USNM). 1 mi NE of Belmont on Rt. 82, 2281 m, July 13, 1980, R. T. Schuh and G. M. Stonedahl, Ephedra sp. (Ephedraceae) , 73, 16♀ (AMNH). Mercury, 19 M, June 23, 1965, H. Knight, J. Merino, Ephedra nevadensis (Ephedraceae) , 3♀ (USNM). Mercury, TM, June 14, 1965, Beck, Knight, Merino, Ephedra nevadensis (Ephedraceae) , 9♀ (USNM); holotype: male (USNM). Utah: Garfield Co.: 14.3 mi S of Rt. 95 on Rt. 276 (2.4 mi N of Star Springs turnoff), 5000 ft, June 19, 1983, R. T. Schuh and M. D. Schwartz, 13 (AMNH). Capitol Reef Natl. Park, Grand Wash­Cohab Canyon Trail, 5350­6640 ft, June 21, 1983, R. T. Schuh and M. D. Schwartz, Ephedra viridis (Ephedraceae) , 23, 4♀ (AMNH). Washington Co.: Rt 15 about 10 mi W of I­15 toward Zion Natl. Park, 1095 m, May 18, 1978, R. T. Schuh, Ephedra sp. (Ephedraceae) , 3♀ (AMNH).

Megalopsallus ephedrellus , new species

Figures 8 , 13

HOLOTYPE: Male, Texas: Crockett Co.: 16.7 mi west Ozona , May 9, 1997, Gillogly & Schaffner. Deposited in the American Museum of Natural History.

DIAGNOSIS: Recognized by pale orange coloration of head, thorax, and hemelytra anterior to apex of scutellum, and dirty greenish coloration posterior to that point; abdomen bright pale green. Size and form of sexual dimorphism similar to nuperus , but that species never with greenish on the hemelytra. Breeds on Ephedra .

DESCRIPTION: Male: Moderately small, total length 2.97–3.25, length apex clypeus–cuneal fracture 2.05–2.24, width across pronotum 0.98–1.07. COLORATION: Head, thorax, appendages, and hemelytra anterior to apex of scutellum pale orange, remainder of hemelytra dirty pale green (fig. 8); abdomen bright pale green; eyes red. SURFACE AND VESTITURE: Dorsum smooth, dull, or very weakly shining, clothed with dark, recumbent, simple setae intermixed with silvery, weakly flattend, somewhat woolly setae. STRUCTURE: Relatively stout­bodied, corial margins nearly straight (fig. 8); labium short, just surpassing posterior margin of fore trochanters; claws elongate, smoothly curving, pulvilli minute. MALE GENITALIA: Vesica relatively large, forming a weak coil, apex relatively strongly attenuated and extending well past secondary gonopore, gonopore subtended by a well­developed gonopore sclerite (fig. 13).

Female: Total length 2.64–2.78, length apex clypeus–cuneal fracture 1.94 –2.04, width across pronotum 0.96–1.01; more strongly ovoid than male, hemelytra nearly conforming to outline of abdomen (fig. 8).

ETYMOLOGY: Named for the host genus Ephedra .

HOST: Ephedra sp. (Ephedraceae) .

DISTRIBUTION: Known only from the type locality near Ozona in western Texas .

DISCUSSION: This is one of four species currently placed in Megalopsallus that is recorded as feeding on Ephedra . The bifid apex of the vesica suggests that the other three species are more closely related to one another than any one of them is to ephedrellus .

PARATYPES: Same data as holotype: 6 males, 52 females ( AMNH; TAMU; USNM) .

Megalopsallus femoralis Kelton

Figures 7 , 13

Megalopsallus femoralis Kelton, 1980: 285 (n. sp.).

DIAGNOSIS: Recognized by somber coloration and relatively small size. Similar in size and general appearance to marmoratus , but lacking marmorate membrane. Similar in coloration to brittoni but smaller.

REDESCRIPTION: Male: Moderately small, total length 3.09–3.24, length apex clypeus– cuneal fracture 2.16–2.29, width across pronotum 0.99–1.07. COLORATION: General coloration somber, head, pronotum, scutellum, eyes, legs, and most of undersurface of body brown or reddish brown, remainder of dorsum and most of antennae tan, head and pronotum sometimes with some red spots (fig. 7). SURFACE AND VESTITURE: Body surface dull, never shining; dorsum with brown, recumbent, simple setae intermixed with subappressed, shining, weakly flattened setae. STRUCTURE: Moderately elongate; labium reaching posterior margin of hind trochanters; claws noticeably elongate, curving, pulvilli small. MALE GENI­ TALIA: Vesica S­shaped with a short apical attenuation; gonopore subapical; gonopore sclerite not developed (fig. 13).

Female: Total length 3.05–3.34, length apex clypeus–cuneal fracture 2.24 –2.36, width across pronotum 1.01–1.09; more broadly ovoid, frons conspicuously more protuberant and clypeus more obviously visible from above than in male (fig. 7).

HOST: Salicornia rubra (Chenopodiaceae) .

DISTRIBUTION: Western Great Plains from Alberta south to Colorado.

DISCUSSION: This species is similar to brittoni , especially in coloration and the weak sexual dimorphism. However, available specimens of femoralis are all smaller than those of brittoni , with no size overlap between the two taxa. The male genitalia, although similar, appear to show consistent differences. Also, as here construed, the distributions of the two taxa are nonoverlapping.

SPECIMENS EXAMINED: CANADA. — Alberta: Brooks , August 20, 1957, A. R . and J. E. Brooks, Paratypes: 13 ( CNC). Lethbridge , June 18, 1939, R . W. Salt, 1♀ ( CNC). Manitoba: Red Deer River, August 3, 1937, C. L. Johnston, 13 ( KU). Saskatchewan: Dana , July 25, 1959, A. R . Brooks, Paratypes: 53, ( CNC). Elbow , June 3–June 26, 1960, A. R . Brooks, Paratypes: 113 ( CNC). Elstow , July 11, 1951 – August 29, 1953, A. R . Brooks, L. A. Konotopetz, Salicornia rubra (Chenopodiaceae) , Holotype and Paratypes: 173, 14♀ ( CNC). USA. — Colorado: Jackson Co.: 7 mi NE of Rand, August 16, 1969, J. C. Schaffner, 13 (TAMU). South Dakota: Campbell Co.: Mound City , July 16, 1938, H. C. Severin, 13, 4♀ ( USNM). Wyoming: Carbon Co. : 8 mi W of Rawlins, September 26, 1966, C. W. O’Brien, 13, 2♀ ( UCB). Lincoln Co. : 4 mi NW of Frontier, June 26, 1966, W. Gagne, J. Haddock, 13 ( UCB) .

Megalopsallus flammeus , new species

Figures 2 , 7 , 14

HOLOTYPE: Male, Nevada: Nye Co., 5 mi. W of Berlin Ichthyosaur St. Monument on Rt. 844, el. 6350 feet, July 1, 1983, R. T. Schuh, M. D. Schwartz, Sarcobatus vermiculatus (Hook.) Torr. var. baileyi (Cov.) Jeps. (Chenopodiaceae) . Deposited in the American Museum of Natural History.

DIAGNOSIS: Recognized unequivocally by bright red coloration of the body and appendages; sexual dimorphism strong (fig. 8).

DESCRIPTION: Male: Moderately small, total length 3.08–3.34, length apex clypeus–cuneal fracture 2.02–2.08, width across pronotum 0.96–1.01. COLORATION: General coloration bright red (fig. 7), including legs (except tarsi); pronotum, scutellum, and underside often tending toward black; eyes blackish; membrane smoky with red veins; tibial spines black but without contrasting dark bases. SURFACE AND VESTITURE: Body surface smooth, weakly shining; dorsum with recumbent, brown, simple setae in­ termixed with silvery woolly setae. STRUC­ TURE: Conspicuously elongate, nearly parallel­sided (figs. 2, 7); labium reaching to hind trochanters; claws relatively short and stout, curving only near apex, pulvilli large, extending to apex of claw, apparently adnate to claw over entire length. MALE GENI­ TALIA: Vesica strongly bent at about midpoint; gonopore large, nearly apical; gonopore sclerite long, well sclerotized (fig. 14).

Female: Total length 2.33–2.59, length apex clypeus–cuneal fracture 1.73–1.90, width across pronotum 0.90–0.98; short, broad, ovoid, in distinct contrast to male (fig. 7).

ETYMOLOGY: Named for its distinctive redorange coloration; from the Latin flammeus , flame colored.

HOSTS: Sarcobatus baileyi , S. vermiculatus (Chenopodiaceae) .

DISTRIBUTION: Central Nevada.

DISCUSSION: I have collected large numbers of specimens of this species on hosts identified as Sarcobatus vermiculatus var. baileyi . Although specimens collected by O’Brien are labeled as occurring on S. vermiculatus , I have never encountered specimens on ‘‘true’’ vermiculatus , and conversely, most records of vermiculatus ­feeding species are not known from var. baileyi . Therefore, host data for the bugs suggest that baileyi represents a distinct plant species.

This species was collected at more than one locality on specimens of Sarcobatus baileyi also inhabited by a mite belonging to the genus Balaustium (Erythraeidae) , which was very similar in appearance and coloration to the females of flammeus . Adults of Balaustium spp. are free­living predators of small arthropods, whereas the larvae are ectoparasites of the same. The relationship, if any, of the mites and the bugs in this particular cases is not known. On the occasion of first collecting this brightly colored bug species my field companions and I dubbed it ‘‘the mite mimic’’ in recognition of the rather strong similarity of appearance of the two taxa.

PARATYPES: USA. — Nevada: Lyon Co.: Weeks, May 29, 1967, C. W. O’Brien, Sarcobatus vermiculatus (Chenopodiaceae) , 33, 6♀ (UCB). Nye Co.: 2 mi E of Tonapah, June 7, 1966, W. Gagne, 13, (UCB). 35 mi N of Tonapah, Coyote Hole Spring/Sevier Reservoir, T8 R42E S11 & 23, 6000 ft, June 30, 1983, Schuh and Schwartz, Sarcobatus vermiculatus var. baileyi (Chenopodiaceae) , 353, 142♀ (AMNH, USNM). 5 mi W of Berlin Ichthyosaur State Monument on Rt. 844, 6350 ft, July 1, 1983, R. T. Schuh and M. D. Schwartz, Sarcobatus vermiculatus var. baileyi (Chenopodiaceae) , 353, 83♀ (AMNH, USNM). Northumberland Canyon Rd, Toquima Mts, T14N R44E Sec. 31, 6400 ft, June 28, 1983, R. T. Schuh and M. D. Schwartz, Sarcobatus vermiculatus var. baileyi (Chenopodiaceae) , 73, 28♀ (AMNH). Washoe Co.: Verdi, July 9, 1967, Gagne, 1♀ (UCB).

Megalopsallus froeschneri (Schuh) , new combination Figures 8 , 14

Merinocapsus froeschneri Schuh, 1986: 220 (n. sp.).

DIAGNOSIS: Recognized by nearly black, weakly to strongly shining head, pronotum, and scutellum, these always contrasting with hemelytra, the latter being either pale with red­orange cuneus or entirely red orange. Coloration of dorsum similar to ephedrae , but femora in that species always entirely dark, whereas in froeschneri femora orange at least on distal one­half. The head, pronotum, and scutellum in froeschneri usually more strongly polished and shining than in ephedrae .

REDESCRIPTION: Male: Medium sized, total length 3.09–3.37, length apex clypeus–cuneal fracture 2.16–2.31, width across pronotum 1.02–1.08. COLORATION: Head, pronotum, and scutellum castaneous to nearly black; hemelytra ranging from orange to intensely red­orange, cuneus always orange, membrane, including most of veins, smoky (fig. 8); eyes black; underside of body castaneous; antennae dark; coxae dark, femora dark proximally, orange distally; tibiae pale, tibial spines dark without dark bases. SUR­ FACE AND VESTITURE: Head, pronotum, and scutellum smooth, polished and shining, clothed with recumbent, dark, simple setae intermixed with woolly, silvery setae. STRUCTURE: Relatively broad­bodied, parallel­sided (fig. 8); labium reaching to near apex of middle coxae; claws elongate, smoothly curving, pulvilli small. MALE GENITALIA: Vesica of moderate length, Sshaped, apex bifid with short projections, secondary gonopore removed from apex by distance equal to length of gonopore, gonopore sclerite very small (fig. 14).

Female: Total length 2.98–3.20, length apex clypeus–cuneal fracture 2.10 –2.35, width across pronotum 1.05–1.09; very similar in appearance and coloration to males, nearly parallel­sided (fig. 9).

HOSTS: Ephedra cutleri , E. nevadensis , E. torreyana , E. viridis (Ephedraceae) .

DISTRIBUTION: Mojave Desert in southern California, Nevada, and Utah; New Mexico.

DISCUSSION: Based on the bifid apex of the vesica, froschneri would appear to be most closely related to ephedrae and pallipes , and on coloration most closely related to ephedrae .

SPECIMENS EXAMINED: USA. — California: Inyo Co. : 2 mi E of Westgard Pass summit, White Mountains, 2125 m, July 2, 1980, R. T. Schuh, Paratypes: 73, 12♀ (AMNH); holotype: male (AMNH). Kern Co.: 20 mi NW of Mojave on Hiway 58, June 13, 1983, G. M. Stonedahl, Ephedra nevadensis (Ephedraceae) , Paratypes: 13, 4♀ (AMNH). Los Angeles Co.: Tehachapi Pass, June 6, 1929, R. L. Usinger, 5♀ (UCB). Little Rock, Mojave Desert, May 20, 1937, E. P. Van Duzee, Paratypes: 13 (CAS). Mint Canyon, May 17, 1937, E. P. Van Duzee, Astragalus sp. 1 ♀ (CAS). Mono Co.: Benton Hot Springs, June 8, 1966, W. Gagne, 3♀ (UCB). San Bernardino Co.: Yucca Valley, 6.3 mi N on Old Woman Springs Rd, May 13, 1978, J. D. Pinto, Ephedra sp. (Ephedraceae) , Paratypes: 13, 1♀ (UCR). 10 mi N of Yucca Valley, May 28, 1975, J. D. Pinto, Ephedra sp. (Ephedraceae) , 1♀ (UCR). Providence Mountains State Recreation Area, 4300 ft, May 18, 1982, M. D. Schwartz, Ephedra viridis (Ephedraceae) , Paratypes: 33 (AMNH). Nevada: Lincoln Co.: 5 mi NE of jct. Rts 38 and 93, 2500 ft, May 19, 1982, M. D. Schwartz, Ephedra nevadensis (Ephedraceae) , Paratypes: 33, (AMNH). Nye Co.: Atomic Test Site, Mercury Hiway at Angle Road (A3), 3800 ft, June 8, 1983, Schuh, Schwartz, Stonedahl, Ephedra nevadensis (Ephedraceae) , Paratypes: 113, 28♀ (AMNH). Atomic Test Site, 1mi S of Cane Springs Road at GS 250 (A5), 4100 ft, June 8, 1983, Schuh, Schwartz, Stonedahl, Ephedra nevadensis (Ephedraceae) , Paratypes: 13, 1♀ (AMNH). Mercury, 18 M, June 7, 1965, E. Beck, J. Merino, Ephedra nevadensis (Ephedraceae) , 1♀ (USNM). Mercury, 40M, June 20, 1965, H. Knight, J. Merino, Ephedra nevadensis (Ephedraceae) , 3♀ (USNM). Mercury, 6M, June 15, 1965, H. Knight, J. Merino, Ephedra nevadensis (Ephedraceae) , 2♀ (USNM). Mercury, CM, June 13, 1965, E. Beck, H. Knight, J. Merino, Ephedra nevadensis (Ephedraceae) , 2♀ (CNC, USNM). Mercury, CU, June 13, 1965, Beck, Knight, Merino, 4♀ (USNM). Mercury, TM, June 14, 1965, E. Beck, H. Knight, J. Merino, Ephedra nevadensis (Ephedraceae) , 3♀ (USNM). Northumberland Canyon Rd, Toquima Mts, T14N R44E Sec. 31, 6400 ft, June 28, 1983, R. T. Schuh and M. D. Schwartz, Ephedra sp. (Ephedraceae) , Paratypes: 43, 6♀ (AMNH). 5 mi E of Gabbs off Rt. 844, 5800 ft, July 1, 1983, R. T. Schuh and M. D. Schwartz, Ephedra sp. (Ephedraceae) , Paratypes: 13, 6♀ (AMNH). New Mexico: San Juan Co.: Angel Peak Campground, June 4, 1977, Hanson and Knowlton, 13, 2♀ (CNC). Utah: Emery Co.: 2.4 mi W of Rt. 24 on Goblin Valley Rd, 5500 ft, June 19, 1983, R. T. Schuh and M. D. Schwartz, Ephedra cutleri (Ephedraceae) , Paratypes: 13 (AMNH). Grand Co.: 11 mi SE of jct. Rts 313 & 163 toward Dead Horse Point, 5200 ft, June 11, 1982, M. D. Schwartz, Ephedra viridis (Ephedraceae) , Paratypes: 133, 35♀ (AMNH). San Juan Co.: Goosenecks Overlook, 5000 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, Ephedra torreyana (Ephedraceae) , Paratypes: 43, 1♀ (AMNH). Rt. 63 at Arizona bor­ der, Monument Valley, 5200 ft, June 16, 1983, R. T. Schuh and M. D. Schwartz, Ephedra cutleri (Ephedraceae) , Paratypes: 133, 53♀ (AMNH). Grand Flat near Collins Canyon, 5600 ft, May 28, 1978, D. A. and J. T. Polhemus, Paratypes: 43, 5♀ (JTP). Glen Canyon Recreation Area, 12 mi S of Rt. 263, T40S R14E, 4300 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, Paratypes: 13 (AMNH). 7.7 mi N of Mexican Hat on Rt 261, T41S R18E, 5000 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, Ephedra sp. (Ephedraceae) , 1♀ (AMNH). 2.7 mi W of Rt. 95 on Rt. 263, T37S R17E, 6000 ft, June 18, 1983, R. T. Schuh and M. D. Schwartz, Ephedra torreyana (Ephedraceae) , Paratypes: 23 (AMNH). Head of Lake Canyon near Nokai Dome Road, 4200 ft, May 29, 1978, D. A. and J. T. Polhemus, Paratypes: 13, 1♀ (JTP). Washington Co.: 2 mi NW of Toquerville on Rt 17, 3800 ft, May 25, 1981, M. D. Schwartz, Ephedra viridis (Ephedraceae) , 1♀ (AMNH).

Megalopsallus humeralis (Van Duzee)

Figures 3 , 9 , 14

Sthenarus humeralis Van Duzee, 1923: 162 (n. sp.).

Europiella humeralis: Knight, 1968: 41 (n. comb., host).

Megalopsallus humeralis: Schuh et al., 1995: 389 (n. comb.).

Megalopsallus arizonae Knight, 1968: 45 (n. sp.). NEW SYNONYMY.

Europiella balli Knight, 1968: 44 (n. sp.). NEW SYNONYMY.

Megalopsallus balli: Schuh et al., 1995: 389 (n. comb.).

Europiella brevicornis Knight, 1968: 45 (n. sp.). NEW SYNONYMY.

Megalopsallus brevicornis: Schuh et al., 1995: 389 (n. comb.).

Europiella lycii Knight, 1968: 40 (n. sp.). NEW SYNONYMY.

Megalopsallus lycii: Schuh et al., 1995: 389 (n. comb.).

Europiella rufiventris Knight, 1968: 42 (n. sp.). NEW SYNONYMY.

Megalopsallus rufiventris: Schuh et al., 1995: 389 (n. comb.).

Europiella viridiventris Knight, 1968: 42 (n. sp.). NEW SYNONYMY.

Megalopsallus viridiventris: Schuh et al., 1995: 389 (n. comb.).

DIAGNOSIS: Recognized by frequently dark coloration of most of dorsum and femora, or at least dark coloration of head, pronotum, scutellum, and femora, with pale hemelytra. Most easily confused with nigrofemoratus on basis of size, coloration, and type of sexual dimorphism; separated most easily by differences in structure of male genitalia, vesica being more elongate and slender in nigrofemoratus than in humeralis , and fact that nigrofemoratus feeds on Atriplex and Grayia (Chenopodiaceae) rather than Lycium (Sola­ naceae), the latter appearing as exclusive host of humeralis . Pale specimens separated from Lycium ­feeding nicholi by cuneus being unicolorous with corium, rather than of at least moderately contrasting coloration as in nicholi .

REDESCRIPTION: Male: Small to medium sized, total length 2.57–3.63, length apex clypeus–cuneal fracture 1.74 –2.31, width across pronotum 0.94–1.11. COLORATION: Dorsum varying from largely deep brown to nearly entirely pale brown, often with head, pronotum, and scutellum dark, hemelytra somewhat lighter to much lighter (fig. 9); eyes blackish; underside mostly dark in dark specimens, abdomen often pale green in lighter specimens; antennae usually reddish, sometimes almost entirely pale, or brownish; femora dark or darkened, tibiae pale, tibial spines dark with dark bases. SURFACE AND VESTITURE: Dorsum smooth, very weakly shining, clothed with recumbent, dark, simple setae intermixed with woolly, silvery setae (figs. 3, 9). STRUCTURE: Weakly to strongly elongate, parallel­sided; specimens in some populations longer than those in others (fig. 9); head strongly declivent (fig. 3); labium reaching onto, but not beyond, hind trochanters; claws relatively short and straight, curving sharply near apex; pulvilli large, adnate to nearly entire ventral surface of claw. MALE GENITALIA: Vesica forming a weak coil, apex not attenuated, gonopore apical; gonopore sclerite well developed (fig. 14).

Female: Total length 2.48–3.30, length apex clypeus–cuneal fracture 1.83–2.19, width across pronotum 0.97–1.16; ovoid; not showing the nonoverlapping size variation seen in males (fig. 9).

HOSTS: Lycium andersonii , L. berlandieri , L. b. parviflorus, L. cooperi , L. fremontii , L. parishii , L. sp. ( Solanaceae ). Probable sitting records: Ambrosia dumosa (Asteraceae) ; Larrea divaricata (Zygophyllaceae) ; Prosopis juliflora (Fabaceae) ; Rhamnus sp. (Rhamnaceae) . Frequently collected when the plants are in fruit.

DISTRIBUTION: Sonora and Baja California, Mexico north as far as southern Utah in interior western North America .

DISCUSSION: Knight (1968) described the new species arizonae , balli , brevicornis , lycii , rufiventris , and viridiventris from the southwestern United States. Comparison of the types and other specimens examined by Knight with holotype of Megalopsallus humeralis (Van Duzee) [‘‘Loreto, Baja California, May 20, 1921, EP Van Duzee Collec­ tor,’’ male (CAS)], including dissections of the male genitalia of most of these nominal species, indicates that even though there is substantial variation in color and size, the male genitalia are very similar for all of them and distinct from all other species of Megalopsallus . I am therefore treating all of these nominal taxa as synonymous, humeralis having priority. The holotype and paratypes of humeralis are faded, like many other Van Duzee specimens, appearing somewhat lighter in color than would be expected of fresh material.

Megalopsallus humeralis shows substantial size variation. The structure of the male genitalia is relatively constant across the range of other forms of variation. Table 1 presents the measurements for this species in two nonoverlapping groupings of ‘‘small’’ and ‘‘large’’ specimens, reflecting the fact that in most populations the specimens are either relatively large or relatively small.

A massive amount of material with carefully documented hosts indicates that humeralis invariably feeds on Lycium spp. The records of specimens from other plant groups offer little evidence for them as alternative hosts. I have collected extensively on all of the plant taxa listed above under ‘‘hosts,’’ and never recorded Megalopsallus breeding on any other than Lycium .

SPECIMENS EXAMINED: MEXICO. — Baja California Norte: 36 mi SE of El Rosario, Rancho El Progresso, March 25, 1979, J. D. Pinto, Lycium sp. (Solanaceae) , 13, 4♀ (UCR). Baja California Sur: Loreto, May 20, 1921, E. P. Van Duzee, Paratypes: 23, 6♀ (CAS). Sonora: 40 mi W of Moctezuma, April 27, 1981, D. A. and J. T. Polhemus, Rhamnus sp. (Rhamnaceae) , 1♀ (JTP). USA. — Arizona: Aztec, February 3, 1941 – April 4, 1942, L. L. Stitt, 4♀ (USNM). Cochise Co. : Apache Trail, March 27, 1951, B. P. Bliven, 13, 4♀ (CAS). Gila Co. : 5.5 mi W of Roosevelt Dam at Apache Lake, 1750 ft, May 27, 1983, Schuh, Stonedahl, and Massie, 2♀ (AMNH). Salt River Mts, 1300 ft, May 9, 1926, A. A. Nichol, 13 (USNM). Maricopa Co. : Gila Bend , 260 m, May 7, 1978, R. T. Schuh, 13 (AMNH). Usery Mtn. Park, January 17, 1983, J. T. Polhemus, 153, 15♀ (JTP). Tempe, April 8, 1940, L. L. Stitt, Paratypes: 13, 2♀ (USNM). 10 mi S of Sun­ flower, April 18, 1982, D. A. and J. T. Polhemus, 23, 1♀ (JTP). Sand Tank Mountains, SE of Gila Bend , 935 m, May 8, 1978, R. T. Schuh and A. F. Guenther, Larrea divaricata (Zygophyllaceae) , 1♀ (AMNH). Sand Tank Mountains, SE of Gila Bend , 900–1000 m, March 24, 1981, R. T. Schuh, Lycium sp. (Solanaceae) , 23, 7♀ (AMNH). Sand Tank Mountains, SE of Gila Bend , 900–1000 m, March 24, 1981, R. T. Schuh, Lycium parishii (Solanaceae) , 273, 54♀ (AMNH). Salt River Canyon at Apache Lake, 2000 ft, April 28, 1981, J. T. Polhemus, 13 (JTP). Phoenix, Arizona Canal at McDowell Rd, April 19, 1982, D. A. and J. T. Polhemus, 13, 7♀ (JTP). Gila Bend , February 3, 1941 – June 1, 1941, L. L. Stitt, Paratypes: 63, 7♀ (USNM). Woolsey Wash near Painted Rock Dam, 205 m, April 3, 1981, R. T. Schuh, Lycium fremontii (Solanaceae) , 83, 12♀ (AMNH). 14 mi S of Wickenburg on Vulture Mine Road, 600–700 m, April 1, 1981, R. T. Schuh and M. D. Schwartz, Lycium andersonii berlandieri (Solanaceae) , 693, 68♀ (AMNH). 5 mi S of Freeman SE of Gila Bend , 625 m, May 8, 1978, R. T. Schuh and A. F. Guenther, Lycium sp. (Solanaceae) , 313, 75♀ (AMNH). 14 mi S of Wickenburg on Vulture Mine Road, 600–700 m, April 1, 1981, R. T. Schuh and M. D. Schwartz, Lycium sp. nr. parishii (Solanaceae) , 213, 11♀ (AMNH). 14 mi S of Wickenburg on Vulture Mine Road, 600–700 m, April 1, 1981, R. T. Schuh and M. D. Schwartz, Lycium sp. (Solanaceae) , 63, 6♀ (AMNH). 16 mi E of milepost 22 on Rt. 85, Sauceda Mts, 565 m, April 2, 1981, R. T. Schuh, Lycium sp. (Solanaceae) , 233, 30♀ (AMNH). 24 mi E of Gila Bend, Freeman , 530 m, May 8, 1978, R. T. Schuh and A. F. Guenther, 13, 3♀ (AMNH). 4 mi S of Wickenburg on Vulture Mine Road, 700 m, April 1, 1981, R. T. Schuh and M. D. Schwartz, Lycium sp. nr. parishii (Solanaceae) , 93, 22♀ (AMNH). 5 mi N of Wickenburg on Rt 93, April 29, 1981, D. A. and J. T. Polhemus, 1♀ (JTP). 5 mi S of Freeman SE of Gila Bend , 575 m, March 24, 1981, R. T. Schuh, Lycium parishii (Solanaceae) , 133, 32♀ (AMNH). 1.8 mi W of Roosevelt on Rt. 88, May 10, 1980, J. D. Pinto, Lycium sp. (Solanaceae) , 53, 9♀ (UCR). Mojave Co.: 9 mi W of Beaver Creek and Virgin River on US 15, Cedar Pocket Rest Stop, 2600 ft, May 12, 1982, M. D. Schwartz, Ambrosia dumosa (Asteraceae) , 3♀ (AMNH). Pima Co. : Organ Pipe Cactus Natl. Mon., Quitobaquito, April 3, 1966, C. W. O’Brien, 23, 4♀ (UCB). Rillito, March 8, 1988, W. A. Jones, Lycium fremontii (Solanaceae) , 33, 5♀ (USNM). Tucson, Santa Catalina Mts, Bear Creek near Saddleback Dr., February 24, 1995, M. D. Schwartz, Lycium berlandieri (Solanaceae) , 103, 11♀ (AMNH). Tucson, April 19, 1926, A. A. Nichol, Paratypes: 23, 4♀ (USNM). Tortolito Mts, 2500–3000 ft, April 22, 1982, M. D. Schwartz, Prosopis juliflora (Fabaceae) , 13 (AMNH). Santa Catalina Mts, Finger Rock Canyon Trail , 3000–3500 ft, April 5, 1981, M. D. Schwartz, Lycium sp. (Solanaceae) , 73, 19♀ (AMNH). Organ Pipe Cactus Natl. Mon., Alamo Wash, 700–800 m, March 26, 1981, R. T. Schuh, M. D. Schwartz, Lycium sp. (Solanaceae) , 453, 59♀ (AMNH). Organ Pipe Cactus Natl. Mon., Alamo Wash, 700– 800 m, March 26, 1981, R. T. Schuh, M. D. Schwartz, Lycium parishii (Solanaceae) , 343, 80♀ (AMNH). Marana, March 9, 1988, W. A. Jones, Lycium sp. (Solanaceae) , 53, 2♀ (USNM). 7 mi W of Tucson, March 9, 1988, W. A. Jones, 23, 4♀ (USNM). 14 mi S of Ajo, April 2, 1966, L. & C. W. O’Brien, 13 (UCB). Organ Pipe Cactus Natl. Mon., Alamo Wash, 700–800 m, R. T. Schuh, M. D. Schwartz, Lycium berlandieri parviflorus (Solanaceae) , 713, 94♀ (AMNH). Pinal Co.: Sacaton, January 22, 1936, L. L. Stitt, Lycium sp. (Solanaceae) , Paratype: 13, (USNM). Yavapai Co.: Coldwater, April 6, 1940, L. L. Stitt, Paratypes: 53, 6♀ (USNM). Castle Wash N of Phoenix, March 21, 1980, J. T. Polhemus, 6♀ (JTP). Yuma Co. : December 20, 1939, L. L. Stitt, Paratypes: 13, 1♀ (USNM). California: Imperial Co. : 1 mi E of Coyote Wells, April 23, 1980, Schwartz and Russell, 1♀ (AMNH). Kern Co.: 7 mi N of Mojave, April 23, 1966, C. W. O’Brien, 23, 4♀ (UCB). Kings Co.: 8 mi SE of Avenal, April 16, 1966, C. W. O’Brien, 13 (UCB). Los Angeles Co.: Palmdale, April 14, 1932, E. P. Van Duzee, 143, 6♀ (CAS). Riverside Co.: Andreas Canyon, Palm Springs, March 11, 1955, W. R. M. Mason, 13 (CNC). San Bernardino Co.: 1.3 mi S of Goffs, 845 m, May 16, 1978, R. T. Schuh, Lycium sp. (Solanaceae) , 563, 64♀ (AMNH). 10 mi W of Lucerne Valley, 905 m, May 13, 1978, R. T. Schuh and J. D. Pinto, Lycium sp. (Solanaceae) , 103, 16♀ (AMNH). 26.7 mi S of Barstow on Rt. 247, 1000 m, May 2, 1985, R. T. Schuh and B. M. Massie, Lycium cooperi (Solanaceae) , 33 (AMNH). Nevada: Clark Co.: 5.8 mi W of Valley of Fire State Park, 845 m, May 17, 1978, R. T. Schuh, 83, 8♀ (AMNH). Nye Co.: Atomic Test Site, Jackass Flats Road, 3300 ft, June 6, 1983, Schuh, Schwartz, and Stonedahl, Lycium andersonii (Solanaceae) , 383, 36♀ (AMNH). Mercury, CM, June 12, 1965 – June 13, 1965, Knight and Merino, 3♀ (USNM). Atomic Test Site, 2.6 mi W of Mercury Hwy on Cane Springs Road, 3400 ft, June 6, 1983, Schuh, Schwartz, and Stonedahl, Lycium andersonii (Solanaceae) , 43, 5♀ (AMNH). Mercury, 6M, June 15, 1965, Knight and Merino, 33 (USNM). Atomic Test Site, Tweezer Rd at Orange Blossom Rd, 4750 ft, June 8, 1983, Schuh, Schwartz, and Stonedahl, Lycium andersonii (Solanaceae) , 1♀ (AMNH). Utah: San Juan Co.: 7.7 mi N of Mexican Hat on Rt. 261, 5000 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, Lycium andersonii (Solanaceae) , 13, 1♀ (AMNH). The Goosenecks Overlook, 5000 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, Lycium andersonii (Solanaceae) , 13, 1♀ (AMNH).

Megalopsallus knowltoni (Knight)

Figures 4 , 9 , 14

Figures

Fig. 7. Habitus photographs of Megalopsallus spp. Note that locality data in this and subsequent figures are associated with specimens by moving from left to right, beginning in row 1 and working down the page. M. atriplicis (3: Nevada, Nye Co., 2.5 mi NE of Gabbs, Rifle Range. ♀: Texas, Cameron Co, Brownsville). M. brendae (3: California, San Bernardino Co., 1.3 mi S of Goffs. ♀: Nevada, Clark Co., west entrance to Valley of Fire State Park). M. brittoni (3 and ♀: Texas, Nueces Co, Corpus Christi). M. californicus (3 and ♀: California, Kern Co., Schafter). M. femoralis (3 and ♀: Canada, Saskatchewan, Elstow). M. flammeus (3 and ♀: Nevada, Nye Co., 35 mi N of Tonapah, Coyote Hole Spring/Sevier Reservoir).

Fig. 13. Male genitalic structures, vesicae of Megalopsallus spp. gs = gonopore sclerite.

Fig. 8. Habitus photographs of Megalopsallus spp. M. froeschneri (3: Nevada, Nye Co., Northumberland Canyon Road, Topquima Mountains, T14N R44E Sec 31. ♀: Utah, San Juan Co., Rt. 63 at Arizona border, Monument Valley; California, San Bernardino Co., Providence Mts State Rec. Area; same as female no. 1). M. ephedrae (3: Nevada, Nye Co., 15.5 mi E of Rt. 376 on Northumberland Mine Rd, T31N R45E Secs 3, 4, 10; Nevada, Lincoln Co., 5 mi NE of jct. Rts 38 & 93; Mexico, Baja California Norte, 12 mi E of El Rosario. ♀: Nevada, Lincoln Co., 5 mi NE of jct. Rts 38 & 93; California, San Bernardino Co., Providence Mts State Rec. Area; California, Inyo Co., 2 mi E of Westgard Pass Summit, White Mts). M. ephedrellus (3 and ♀: Texas, Crockett Co., 16.7 mi W of Ozona).

Fig. 14. Male genitalic structures, vesicae of Megalopsallus spp. Left paramere and phallotheca of M. knowltoni. gs = gonopore sclerite.

Fig. 2. Habitus of Megalopsallus flammeus, male (Nevada, Nye Co., 35 mi N of Tonapah, Coyote Hole Spring/Sevier Reservoir).

Fig. 3. Megalopsallus humeralis, male, scanning micrographs. A. Lateral view of head and thorax. B. Mesothoracic spiracle and metathoracic scent gland evaporatory area. C. Detail of setae on thoracic pleuron.

Fig. 9. Habitus photographs of Megalopsallus spp. M. humeralis (3: Arizona, Pima Co., Santa Catalina Mts, Finger Rock Canyon Trail. ♀: Nevada, Nye Co., Mercury. 3: Arizona, Pima Co., Santa Catalina Mts, Bear Creek near Saddleback. ♀: California, San Bernardino Co., 10 mi W of Lucerne Valley). M. knowltoni (3 and ♀: California, Mono Co., 8 mi W of Nevada state line on Rt. 359). M. marmoratus (3: Mexico, Baja Calif. Norte, 1 mi S of Bahia de los Angeles. ♀: Arizona, Maricopa Co., just S of Buckeye). M. nicholi (3: Mexico, Zacatecas, 13 mi SW of Concepcion del Oro. ♀: Mexico, Zacatecas, 6 mi S of Concepcion del Oro). M. nigricaput (3 and ♀: Arizona, Graham Co., Stockton Pass, Pinaleno Mts).

Fig. 4. Megalopsallus knowltoni, female, scanning micrographs. A. Lateral view of head and thorax. B. Mesothoracic spiracle and metathoracic scent gland evaporatory area. C. Detail of setae on thoracic pleuron. D. Ventral view of pretarsus.