Chlamydatus Curtis, 1833

SCHUH, RANDALL T. & SCHWARTZ, MICHAEL D., 2005, Review of North American Chlamydatus Curtis Species, with New Synonymy and the Description of Two New Species (Heteroptera: Miridae: Phylinae), American Museum Novitates 3471 (1), pp. 1-56 : 12-20

Chlamydatus Curtis

Chlamydatus Curtis 1833: 198 (n.gen.).

REVISED DIAGNOSIS: Small, elongate oval or ovate, sometimes submacropterous with membrane shortened and just covering abdomen, or brachypterous, with hemelytra truncate, leaving much of abdomen exposed (figs. 1, 2); total length 1.28–3.09, width across pronotum 0.67–1.03; coloration at least partially dark brown or castaneous, often entirely black (figs. 1, 2); eyes somewhat bulging and extending laterally beyond anterolateral angles of pronotum (figs. 7A,B, 8A, 9A,B); posterior margin of vertex round­ ed, sometimes broadly so (figs. 7A, 9A); antennae not sexually dimorphic, segment 2 usually slender, rarely fusiform, generally no longer than width of head across eyes (figs. 1, 2); claws only moderately elongate, nearly straight over much of length, pulvilli relatively large, adnate to at least basal half of ventral claw surface (figs. 7E,F, 8C,D, 9D); parempodia setiform. Vestiture of dorsum composed of simple or weakly woolly setae, these ranging from dull black to silvery or golden and shining, sometimes appearing neatly arranged, less frequently having a somewhat disordered appearance (e.g., C. associatus , C. obliquus (Uhler)) (figs. 1, 2, 7C, 8B, 9D). Femora moderately broad (fig. 7E). Genitalia (figs. 4–6, 7H, 8F, 9F): Vesica with a single strap, apical attenuation single, varying greatly from about the length of the gonopore to about half the length of the vesica, and occasionally with a ‘‘secondary spine’’ on the apical attenuation (see C. opacus , C. schuhi Knight ) or with apical spicules (see C. monilipes , C. montanus Knight ); gonopore relatively large, closed both proximally and distally, distal margin broadly rounded, without area of attenuation, proximally without a gonopore sclerite (as found in Atractotomus Fieber [see Stonedahl, 1990]); phallotheca without distinctive features; left paramere conventionally phyline, without distinctive structural variation; right paramere lanceolate.

DISCUSSION: Dark­colored Chlamydatus spp. will be most easily confused with some of the small black species of Plagiognathus Fieber. The male genitalia in the two groups are absolutely distinctive from one another, however, Chlamydatus having a vesica formed of a single strap with an attenuated apex, whereas the vesica in Plagiognathus has two straps and terminates in two broad blades; on that basis they can be unequivocally separated. Chlamydatus spp. are similar in size and coloration to some other North American taxa, such as species of Arctostaphylocoris Schuh and Schwartz, 2004 and Phoenicocoris spp. (see Schwartz and Stonedahl, 2004), but those taxa always have some scalelike setae on the dorsum and usually on the thoracic pleuron, whereas Chlamydatus spp. never possess scalelike setae.

Chlamydatus is a taxon of longstanding recognition. Nonetheless, producing a straightforward diagnosis for the group on the basis of external and genitalic characters has not proved to be a simple matter. Whereas most specimens of Chlamydatus in reasonably good condition can be recognized without the necessity of dissecting the male genitalia, placing specimens to genus will often be the more difficult part of the identification process. The labium varies considerably in length within the genus. Its use in identification of species can be frustrating because of intraspecific variation, sexual dimorphism, and because of flexion of both the head and the labium. Consequently, we have largely avoided using labial length as a diagnostic feature. We have, however, used total length of specimens as a way of distinguishing many taxa. Experience suggests that attempting to determine size simply by inspection will produce inaccurate results. Thus, we recommend performing all measurements with an ocular micrometer.

Many of the New World species of Chlamydatus for which hosts are known feed on members of the Asteraceae , both herbaceous and woody. Several appear to show substantial host specificity, such as C. becki and C. brevicornis on Artemisia spp. and C. californicus , n. sp., on Hemizonia pungens ; it is these species that are usually least well represented in collections. A few species, such as C. associatus and C. suavis (Reuter) are widespread and appear to feed on a variety of ruderal plants, including Asteraceae , and occasionally on cultivated species. Specimens of these species are often acquired by general collectors through sweeping plants in early successional stages and consequently these are the species most commonly encountered in collections, often without host data.

There is no obvious pattern of host relationships for those taxa that do not feed on Asteraceae . For example, C. obliquus (Uhler) feeds primarily on herbaceous legumes in the genera Lupinus and Astragalus . Chlamydatus uniformis (Uhler) feeds on perennial members of the Lamiaceae in the genus Salvia . Chlamydatus schuhi Knight , although undoubtedly known to breed on Asteraceae , is also known from species of Leptodactylon and Phlox in the Polemoniaceae . Chlamydatus keltoni , n. sp., has been recorded from a variety of plant families, including— among others—the Fabaceae ( Astragalus , Lupinus , and Medicago spp. ), Ranunculaceae ( Ranunculus sp. ), and Rosaceae ( Potentilla spp. ); a similar pattern of host associations is seen for C. pulicarius (Fallén) . Host data appear to be completely lacking for some species, including C. opacus (Zetterstedt) , C. ruficornis Knight , and C. wilkinsoni (Douglas and Scott) .

KEY TO NORTH AMERICAN SPECIES OF CHLAMYDATUS CURTIS

1. Fore­ and middle femora entirely pale yellowish to white, never with basal infuscation, posterior femur dark brown to nearly black on proximal 90%, pale distally, in marked contrast to fore­ and middle femora (fig. 3A)......................... 2

— Coloration of femora not as above (fig. 3A), coloration of hind femora not strongly contrasting with that of fore­ and middle femora (fig. 3B–I)................. 6

2. Large species, length 2.78–2.98; frequently with clavus, exocorium, and posterior margin of pronotum pale, less commonly dorsum entirely black (fig. 1); dorsal vestiture relatively dense, somewhat woolly, shining (fig. 1); often feeding on Lupinus or other herbaceous legumes; interior of western North America....... obliquus (Uhler)

— Smaller species, length less than 2.76; coloration variable; vestiture woolly or neatly arranged........................ 3

3. Hemelytra largely, and sometimes posterior margin of pronotum, pale; scutellum always dark (fig. 2); known from Leptodactylon and Phlox (Polemoniaceae) and Chrysothamnus spp. Asteraceae ); Great Basin................. schuhi Knight

— Dorsum unicolorous dark............. 4

4. Antennal segment 2 longer than width of head across eyes (75:68); body form in males relatively elongate, hemelytra nearly parallel­sided (fig. 1); Ambrosia spp. (Asteraceae) and other annuals; widely distributed in North America................................ associatus (Uhler) — Antennal segment 2 shorter than width of head across eyes (56:69, 59:70); body form broader and hemelytra convexly rounded laterally.................. 5

5. Dorsum clothed with moderately shining setae (fig. 1); labium at most reaching slightly beyond posterior margin of middle trochanters; antennal segment 2 in males usually entirely dark; known only from Artemisia cana ; southeastern Alberta ........................ artemisiae Kelton

— Dorsum clothed with neatly arranged, very weakly shining, dark setae (fig. 2); labium reaching beyond posterior margin of hind trochanters; antennal segment 2 in males usually mostly pale; known from Potentilla ; montane western North America..................... pallidicornis Knight

6. All femora and tibiae completely black (fig. 3F)............................. 7

— All femora and tibiae not completely black; femora either pale on distal 10% (fig. 3G– I), or if unicolorous then femora entirely infuscate or reddish (fig. 3C–E), or femora infuscate or reddish on proximal half only (fig. 3B); tibiae either infuscate (fig. 3C– E) or pale, spines with weakly to strongly darkened bases (fig. 3B,G–I)........ 8

7. Larger species, length 2.65–2.88; jet black, shining; known from Salvia spp. ; coastal southern California and Baja California Norte.............. uniformis (Uhler)

— Smaller species, length 2.21–2.23; brownish black, weakly shining; known from Artemisia spp. ; Mojave Desert and Great Basin....................... becki Knight

8. Hemelytra covering entire abdomen, with at least a limited area of membrane present; very rarely membrane strongly reduced and pygophore exposed in males (fig. 1, opacus )......................... 9

— Hemelytra truncate and membrane completely absent (staphylinoid) (fig. 1, californicus , fig. 2, ruficornis , wilkinsoni ); at least half of abdomen exposed.......... 18

9. Dorsum cream and brown, not unicolorous dark; membrane greatly reduced, exposing pygophore in male (fig. 1); northern Canada............. opacus (Zetterstedt)

— Dorsum unicolorous dark, usually blackish, castaneous, or very deep brown; membrane sometimes reduced, but never exposing pygophore................ 10

10. Femora and tibiae conspicuously reddish, although areas of femora may be much darker than remainder of legs...... 11

— Femora and tibiae never conspicuously reddish; if not black or very dark brown, then at most yellow or yellow orange.... 12

11. Dorsum dull, slate black (fig. 2); antennal segment 2 weakly fusiform, red (fig. 2); frons distinctly bulging (fig. 2); host unknown; plains of North America........................ ruficornis Knight

— Dorsum castaneous, polished and moderately shining (fig. 1); antennal segment 2 linear, usually black in males (fig. 1), sometimes paler in females; frons weakly convex (fig. 1), but never bulging; known from Artemisia spp. ; montane western North America...... montanus Knight

12. All femora mostly pale, ornamented with some black spots (see fig. 3B)..... 13

— All femora largely dark, usually black, pale only on distal one­fourth or less (fig. 3G) .............................. 14

13. Smaller species, total length 2.40–2.41; apical portion of phallotheca tapered, barely extending onto posterior surface of genital capsule; apex of vesica tapering towards apex (fig. 6); membrane in males apparently always fully developed, exceeding apex of abdomen (fig. 2); hosts in New World unknown; Yukon Territory and north­central United States.............................. pullus (Reuter)

— Larger species, total length 2.51–3.09; apical portion of phallotheca greatly elongate and slender, more or less parallel sided, extending well onto posterior surface of genital capsule (figs. 5, 9F); apex of vesica greatly elongated and nearly parallel sided (fig. 5); membrane often reduced, hemelytra often not exceeding apex of abdomen in males and females (fig. 2); from annual Asteraceae and other herbaceous spp.; north of 45°N latitude and further south in Rocky Mountains and scattered localities in eastern North America....................... pulicarius (Fallén)

14. Large species, total length greater than 2.50; distal portion of genital capsule broad, apical portion of phallotheca greatly elongate, but not attenuate distally, strongly extending over posterior surface of genital capsule (figs. 5, 9F); apex of vesica greatly elongated and nearly parallel sided (fig. 5); known from annual Asteraceae and other herbaceous spp.; north of 45°N latitude and further south in Rocky Mountains and scattered localities in eastern North America..... pulicarius (Fallén)

— Smaller species, total length less than 2.60, usually less than 2.50; if distal portion of genital capsule broad with apical portion of phallotheca elongate and extending over posterior surface of genital capsule, then phallotheca sharply attenuate apically (figs. 4, 8F); apex of vesica variable, but never so greatly elongated and parallel sided.......................... 15

15. Tibiae entirely pale, tibial spines without dark bases (fig. 3G); coloration often distinctly brownish rather than blackish; recorded from Ambrosia sp. ; widely distributed................ suavis (Reuter)

— Tibiae pale, tibial spines with distinctly darkened bases (fig. 3H,I); coloration of dorsum variable................. 16

16. Dorsum densely clothed with conspicuously shining, somewhat woolly pubescence (fig. 1); known from Artemisia ; British Columbia , south to northern California, east to Colorado .... brevicornis Knight

— Dorsal vestiture neatly arranged and appearing black when viewed from above, never with a woolly or shaggy appearance.... ............................... 17

17. Hind femur with a very narrow pale band at tibial articulation (fig. 3I); all tibiae white with heavy black spots at bases of spines (fig. 3I); always macropterous, without hemelytral polymorphism as below (fig. 1); vesica as in figure 5; known from Haplopappus and other woody Asteraceae ; Great Basin, Arizona, and California ................... monilipes Van Duzee

— Hind femur more broadly pale at tibial articulation (fig. 3H); tibiae yellowish, spots at bases of tibial spines smaller, brown (fig. 3H); most populations showing hemelytral polymorphism, containing both macropterous and submacropterous individuals, males usually macropterous, females usually submacropterous (fig. 1); vesica as in figure 4; known from Lupinus and Potentilla ; Alaska, western Canada, and Rocky Mountains of the United States................ keltoni , new species

18. Antennal segment 2 weakly fusiform (fig. 2); antennal segments 1 and 2 largely and distinctly reddish (fig. 2); host association unknown; plains of North America........................ ruficornis Knight

— Antennal segment 2 linear; antennal segments 1 and 2 not distinctly reddish.... ............................... 19

19. Hemelytra dark to nearly black (fig. 2); small species, total length 1.28–1.89; host association in New World unknown; Alaska, Canada............................... wilkinsoni (Douglas and Scott)

— Hemelytra brown (fig. 1); larger species, total length 1.60–1.94; known from Hemizonia pungens (Asteraceae) ; central valley of California .................................. californicus , new species

Chlamydatus artemisiae Kelton

Figures 1 , 3A , 4

Chlamydatus artemisiae Kelton, 1965: 1138 (n.sp., host).

DIAGNOSIS: Recognized by the moderate size, total length 2.33–2.44, the unicolorous dark dorsum clothed with shining, weakly woolly setae (fig. 1), and yellowish fore­ and middle femora contrasting with the strongly infuscate hind femora (fig. 3A). Femoral coloration as in C. associatus , C. obliquus , C. pallidicornis , and C. schuhi . Distinguished from C. associatus by the broader, less elongate body form and antennal segment 2 distinctly shorter than width of head across eyes, whereas in C. associatus body form more elongate and antennal segment 2 longer than width of head (table 1). Male genitalia similar to those of C. associatus , the vesica being relatively short with a short apical attenuation (fig. 4). Distinguished from C. obliquus specimens with entirely dark hemelytra by the larger size of that species, the total length of males in C. obliquus always being greater than 2.77, in C. artemisiae total length never greater than 2.44. Antennal segments 1 and 2 in males of C. pallidicornis usually entirely pale, whereas C. artemisiae males with antennal segments 1 and 2 dark; vestiture in C. pallidicornis dark and neatly arranged, not shining and weakly woolly as in C. artemisiae . Easily distinguished from C. schuhi by the invariable presence of contrasting pale markings on the posterior margin of the pronotum and on the hemelytra in that species (fig. 2), whereas C. artemisiae always unicolorous dark on the dorsum (fig. 1).

HOST: Artemisia cana (Asteraceae) .

DISTRIBUTION: Known from southeastern Alberta, Canada.

DISCUSSION: This species was described by Kelton (1965) on the basis of numerous specimens from southeastern Alberta on Artemisia cana . Although we find it remarkable that any species of Chlamydatus with such a common host should have so restricted a distribution, we were unable to find other material that compares in body form, vestiture type, antennal length, and genitalic structure.

SPECIMENS EXAMINED: Holotype: male: ‘‘Walsh, Alta[.,] 28.V.1952 [,] L. A. Konotopetz[;] Artemisia cana , Type No. 8907’’; deposited in the Canadian National Collection of Insects, Ottawa.

Additional specimens: CANADA: Alberta: Onefour, May 14, 1952, A. R. Brooks, Artemisia cana (Asteraceae) , paratype, 13 (CNC). Walsh, May 28, 1952, A. R. Brooks, L. A. Konotopetz, Artemisia cana (Asteraceae) , paratypes, 123, 9♀ (CNC).

Chlamydatus associatus (Uhler)

Figures 1 , 3 , 4 , 7

Agalliastes associatus Uhler, 1872: 419 (n.sp.).

DIAGNOSIS: Recognized by the moderate size, the dorsum unicolorous dark and clothed with shining, weakly woolly setae (fig. 1), the fore­ and middle femora yellowish to white—in marked contrast to the strongly infuscate hind femora (fig. 3A)— and the vesica in the male as in figure 4. Femoral coloration as in C. artemisiae , C. obliquus , C. pallidicornis , and C. schuhi . Distinguished from C. artemisiae by the more elongate body form and length of antennal segment 2 being greater than width of head across eyes, whereas C. artemisiae with antennal segment 2 distinctly shorter than width of head (table 1). Male genitalia similar to those of C. artemisiae , the vesica being relative short with a short apical attenuation (fig. 4). Distinguished from C. obliquus specimens with entirely black hemelytra by the larger size of that species, the total length of males in C. obliquus always being greater than 2.77; in C. associatus total length never greater than 2.76. Antennal segments 1 and 2 in males of C. pallidicornis usually entirely pale, whereas associatus males with antennal segments 1 and 2 dark; vestiture in C. pallidicornis dark and neatly arranged, not shining and weakly woolly as in C. associatus . Easily distinguished from C. schuhi by the invariable presence of contrasting pale markings on the posterior margin of the pronotum and on the hemelytra in that species, whereas C. associatus always unicolorous dark on the dorsum.

HOSTS: Ambrosia spp. ( Knight, 1923, 1941; Blatchley, 1926). Label data in collections examined by us indicate that C. associatus breeds on Ambrosia spp. , but has also been collected on a much wider range of plants, in accordance with the list provided by Kelton (1965).

DISTRIBUTION: Widely distributed in North America, ranging from Alaska to central Mexico, and from the Atlantic to the Pacific coasts.

DISCUSSION: As noted by Kelton (1965), this is one of the most common species of Chlamydatus in collections. Blatchley (1926) and Knight (1923, 1941) recorded C. associatus as breeding on Ambrosia spp. in the eastern United States. Kelton (1965) record­ ed the species from a wide variety of plant taxa, including Ambrosia spp. , Artemisia spp. and other Asteraceae , Astragalus sp. , and Medicago sp. in the Fabaceae , as well as representatives of the Brassicaceae and Chenopodiaceae . The specimens we have examined indicate that C. associatus is frequently collected by sweeping of ruderal vegetation. Furthermore, the label data and wide distribution clearly suggest that C. associatus is not restricted to Ambrosia spp. , in conformity with the data presented by Kelton (1965), although like several other Chlamydatus spp. , the Asteraceae may be the dominant host group. Wheeler (2001) cited sources that record C. associatus as a pollen feeder and as an egg predator.

SPECIMENS EXAMINED: Holotype: male (?): ‘‘Ogden [Utah], Agalliastes associatus Uhler , Type No. 1133 USNM’’; deposited in the National Museum of Natural History , Washington, D.C.

Additional specimens: CANADA: Alberta: Brooks, August 20, 1957, A. R. and J. E. Brooks (CNC). Brooks, Special Crops and Horticulture Research Center, July 17, 1990, M. D. Schwartz, Kochia scoparia (Chenopodiaceae) , 13 (AMNH). Conrad, June 21, 1952, A. R. Brooks (CNC). Drumheller, August 11, 1957, A. R. and J. E. Brooks, 13 (CNC). Empress, June 7, 1957, A. R. Brooks, MacNay (CNC). Kananaskis Highway, June 25, 1973, L. A. Kelton (CNC). Kananaskis Road, July 20, 1974, L. A. Kelton (CNC). Lethbridge, July 5, 1956, O. Peck, Medicago sativa (Fabaceae) (CNC). Manyberries, June 21, 1952, L. A. Konotopetz (CNC). Medicine Hat, August 22, 1930, J. H. Pepper (CNC). Milk River, 25 mi E of, June 28, 1956, O. Peck, Brassica sp. (Brassicaceae) (CNC). Onefour, August 19, 1955, A. R. Brooks (CNC). Orkney District, July 13, 1934, G. F. Manson (CNC). Steveville, August 21, 1957, A. R. and J. E. Brooks (CNC). British Columbia : Barriere, waste lot behind Monte Carlo Motel near Rt 5, August 27, 1993, M. D. Schwartz, Berteroa incana (Brassicaceae) , 43 (AMNH). Cawston, 4 mi E of, July 9, 1959, L. A. Kelton, 13 ( Asteraceae ) (CNC). Cranbrook, July 23, 1959, L. A. Kelton ( Pinaceae ) (CNC). Cranbrook, September 23, 1959, L. A. Kelton, 23 (CNC). Fife, 5 mi E of, June 8, 1959, R. E. Leech, 13 (CNC). Grand Forks, August 7, 1950, H. R. MacCarthy ( Apiaceae ) (CNC). Grand Forks, July 7, 1951, D. A. Arnott, 13 (CNC). Grand Forks, June 8, 1959, E. E. MacDougall (CNC). Grand Forks, 9 mi E of, June 8, 1959, L. A. Kelton, 53 (CNC). Grand Forks, 15 mi W of, June 7, 1959, L. A. Kelton, 23 (CNC). Oliver, April 29, May 15, 23, 27, 29, 1959, L. A. Kelton, R. E. Leech (CNC). Oliver, May 27, 1959, Madge, Kelton and Leech, 53 (CNC). Osoyoos Lake, Haynes Ecology Reserve, May 8, 1994, G. G. E. Scudder, Sisymbrium loeselii , 13 (UBC). Vaseaux Lake, July 9, 1959, L. A. Kelton (CNC). Vaseaux Lake, Oliver, May 28, 29, 1959, L. A. Kelton, 23 (CNC). Wasa, August 7, 1970, L. A. Kelton, 13 (CNC). Westbank, June 30, 1955, Wilson, Wakefield (CNC). Manitoba: Aweme, June 22, 1953, A. R. Brooks, L. A. Kelton (CNC). Boissevain, July 17, 1953, A. R. Brooks, L. A. Kelton (CNC). Brandon, July 19, 1946, P. H. Westdal (CNC). Carberry, July 27, 1953, A. R. Brooks, L. A. Kelton (CNC). Carberry, June 14, 1953, Brooks and Kelton, 13 (CNC). Foxwarren, 2.5 mi NW of, on Rt 16, July 19, 1990, M. D. Schwartz, Capsella bursapastoris (Brassicaceae) , 13 (AMNH). Goodlands, August 28, 1920, R. W. White , 13 (CNC). Goodlands, July 28, 1930, R. M. White (CNC) . Melita, August 23, 1953, A. R. Brooks (CNC). Morden, August 9, 1945, W. R. Allen (CNC). Morris, August 15, 1953, A. R. Brooks, 13 (CNC). Pilot Mound, July 31, 1958, A. R. and J. E. Brooks (CNC). Saint Francois Xavier, June 13, 1953, A. R. Brooks, L. A. Kelton (CNC). Souris, July 27, 1953, A. R. Brooks (CNC). Treesbank Ferry, June 5, 1958, J. F. McAlpine, Thlaspi sp. (Brassicaceae) (CNC). Turtle Mountain, July 17, 1953, A. R. Brooks, L. A. Kelton (CNC). Virden, July 9, 1953, A. R. Brooks, L. A. Kelton (CNC). Virden, July 9, 1953, Brooks and Kelton, Helianthus sp. (Asteraceae) , 13 (CNC). Winkler, August 24, 1949, W. R. Allen (CNC). Winnipeg, June 6, 1953, A. R. Brooks, L. A. Kelton (CNC). Ontario: Ariss, May 27, 1962, L. A. Kelton, Thorpe, Antennaria sp. (Asteraceae) (CNC). Aylmer, July 19, 1955, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Belle River, June 7, 1961, L. A. Kelton, G. Brumpton (CNC). Belleville, June 25, 1956, J. Franz (CNC). Brantford, July 16, 1962, L. A. Kelton, Robinia sp. (Fabaceae) (CNC). Britannia, Ottawa, September 1, 1913, W. Metcalfe, Helianthus sp. (Asteraceae) (CNC). Caledonia, July 12, 1955, L. A. Kelton, Helianthus sp. (Asteraceae) (CNC). Cayuga, June 26, 1961, L. A. Kelton, G. Brumpton (CNC). Chatham, August 4, 1933, W. F. Lindsay (CNC). Clinton, September 7, 1961, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Cobourg, July 11, 1981, D. J. E. Brown (CNC). Collingwood, June 13, 1962, L. A. Kelton, Thorpe (CNC). Constance Bay, September 20, 1961, L. A. Kelton (CNC). Delhi, July 20, 1955, L. A. Kelton (CNC). Eagleville, August 17, 1955, L. A. Kelton, 13 (CAS). Effingham, July 12, 1955, L. A. Kelton (CNC). Eramosa, September 5, 1961, L. A. Kelton, G. Brumpton, Ambrosia sp. (Asteraceae) (CNC). Forestville, September 9, 1963, G. G. E. Scudder (CNC). Fort Erie, July 10, 1955, L. A. Kelton, 23 (CAS, CNC). Gore Bay, July 10, 1961, G. Brumpton (CNC). Guelph, August 31, 1963, G. G. E. Scudder (CNC). Haggersville, July 9, 1962, L. A. Kelton, Thorpe, Ambrosia sp. (Asteraceae) (CNC). Harwich, July 13, 1933, G. M. Stirrett (CNC). Horning, June 15, 1961, L. A. Kelton (CNC). Jordan, August 9, 1961, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Kincardine, May 28, 1962, L. A. Kelton, Thorpe (CNC). Kingsville, May 23, 1962, L. A. Kelton, Thorpe (CNC). Leamington, July 2, 1931, G. S. Walley, Helianthus sp. (Asteraceae) (CNC). Newcastle, September 9, 1963, G. G. E. Scudder (CNC). Niagara Falls, July 7, 1955, L. A. Kelton (CNC). Normandale, June 17, 1931, G. S. Walley (CNC). Oakland, July 17, 1962, L. A. Kelton, Thorpe (CNC). One Sided Lake, June 27, 1960, L. A. Kelton, Whitney (CNC). Ottawa, August 22, 1958 and September 1, 1962, L. A. Kelton (CNC). Point Pelee, September 11, 1961, L. A. Kelton (CNC). Point Ryerse, June 1, 1939, G. E. Shewell (CNC). Presquile Province Park, July 1, 1958, G. G. E. Scudder (CNC). Queenston, June 24, 1961, L. A. Kelton, G. Brumpton (CNC). Rainy River, August 3, 1950, L. A. Kelton, Whitney, Ambrosia sp. (Asteraceae) (CNC). Rosemont, May 30, 1962, L. A. Kelton, Thorpe (CNC). Saint Ann’s, June 9, 1962, L. A. Kelton, Thorpe (CNC). Saint Catharines, September 3, 1961, L. A. Kelton (CNC). Sherkston, September 22, 1961, L. A. Kelton (CNC). Simcoe, July 13, 1955, L. A. Kelton (CNC). Thessalon, July 11, 1965, W. Gagne, 33 (UCB). Tillsonburg, July 20, 1955, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Turkey Point, September 9, 1963, G. G. E. Scudder (CNC). Vienna, July 15, 1961, L. A. Kelton (CNC). Vineland, June 9, 1962, L. A. Kelton, Thorpe, Matricaria sp. (Asteraceae) (CNC). Violet Hill, September 5, 1961, L. A. Kelton, G. Brumpton, Ambrosia sp. (Asteraceae) (CNC). Virgil, July 9, 1955, L. A. Kelton (CNC). Waterford, July 17, 1962, L. A. Kelton, Thorpe (CNC). Willard Lake, 5 mi E of, August 10, 1960, L. A. Kelton, Whitney (CNC). Wheatley, July 18, 1955, L. A. Kelton, Medicago sativa (Fabaceae) (CNC). Prince Edward Co.: August 20, 1952, J. F. Brimley (CNC). Quebec: Choisy, nr Rigaud, August 28, 1956, Lindberg (CNC). Gatineau National Park, jct. Meech Lk. Rd and Gatineau Pkwy., September 14, 1994, M. D. Schwartz, Ambrosia artemisiifolia (Asteraceae) , 33 (AMNH). Gatineau National Park, June 14, 1981, D. J. E. Brown (CNC). Hudson Heights, August 25, 1956, Lindberg (CNC). Ile Jesus, 20 mi NW Montreal, near St. Rose, August 19, 1956, J. D. Lattin, 33 (OSU). Montreal, August 19, 1956, Slater, Davis, et al., 33 (AMNH). Sutton, August 5, 1961, G. Brumpton (CNC). Saskatchewan: Elbow, June 20, 1060, A. R. Brooks (CNC). Estevan, June 15, 1955, A. R. Brooks (CNC). Gascoigne, August 23, 1957, A. R. and J. E. Brooks (CNC). Minton, August 11, 1955, A. R. Brooks (CNC). Minton, February 8, 1955, A. R. Brooks, 13 (CNC). Regina, 186 km N of, off of Rt 11, 3.3 km E of Rt 11, July 10, 1990, M. D. Schwartz, Brassica juncea (Brassicaceae) , 23 (AMNH). Rockglen, August 2, 1955, A. R. Brooks (CNC). Saskatoon, August 9, September 14, 1950, L. A. Konotopetz, Helianthus sp. (Asteraceae) (CNC). Tunstal, August 27, 1957, A. R. AND J. E. Brooks (CNC). Val Marie, August 5, 8, 1955, A. R. Brooks, 23 (CNC). Weyburn, July 7, 1957, L. A. Konotopetz, 13 (CNC). Willow Bunch, July 27, 1955, A. R. Brooks (CNC). Wood Mountain, August 5, 1955, A. R. Brooks (CNC). MEXICO: Michoacan: 13 mi S of Nueva Italia, July 9, 1985, Jones and Schaffner, 13 (TAMU). Nuevo Leon: 37 mi NE of China, July 18, 1954, University of Kansas Mexico Expedition, 13 (KU). USA: Alaska: 32 mi N of Seward Kenai Peninsula, July 1, 1957, G. Byers and party, 13 (KU). Arizona: Apache Co. : Vernon, July 25, 1956, Gerhardt and Butler, 13 (UAZ). Cochise Co. : 8 mi S of Apache , September 18, 1978, J. D. Pinto, 13 (UCR). Benson, July 12, 1952, R. H. Beamer, 13 (KU). Coconino Co.: Flagstaff, Coconino National Forest, August 16, 1967, L. A. Kelton (CNC). Flagstaff, July 14, 1947, L. D. Beamer, 13 (KU). Flagstaff, July 9, 1938, D. Rockefeller, 13 (AMNH). Flagstaff, September 9, 1951, A. T. McClay, 53 (UCD). Flagstaff, NW of, August 13, 1975, J. C. Schaffner, 33 (TAMU). Fredonia, June 3, 1965, G. F. Knowlton, 13 (USU). Fredonia, October 10, 1954, K. Goodarzy and G. F. Knowlton, 103 (KU). Grand Canyon, South Rim, June 23, 1930, R. L. Usinger, 23 (UCB). Jacob Lake, Kaibab National Forest, August 12, 1973, L. A. Kelton, Lupinus sp. (Fabaceae) (CNC). Kaibab National Forest, N of Demotte Campground, August 3, 1983, C. B. Barr, 23 (LSU). Kendrick Park, Coconino National Forest, 7400 ft, July 15, 1968, L. A. Kelton (CNC). Mormon Lake, 7000 ft, July 8, 1956, F. G. Werner, 13 (UAZ). No specific locality, August 13, 1927, R. H. Beamer, 13 (KU). Parks, Kaibab National Forest, August 16, 1967, L. A. Kelton (CNC). Sedona, 3.5 mi S of, on Rt 179, 4200 ft, June 15, 1983, R. T. Schuh and M. D. Schwartz, 13 (AMNH). Sunset Crater Road, 1 mi E of Rt 89A, 7300 ft, June 16, 1983, R. T. Schuh, Artemisia sp. (Asteraceae) , 23 (AMNH). Gila Co. : White River , June 19, 1950, L. D. Beamer, 33 (KU). Navajo Co.: Forest Road 45, 3 mi NE of Highway 260, 6800 ft, August 18, 1998, J. C. Schaffner, 23 (TAMU). Lewis Canyon Campground, Pinedale, 2 mi S of, 6700 ft, May 30, 1983, R. T. Schuh and G. M. Stonedahl, Lupinus sp. (Fabaceae) , 23 (AMNH). Show Low, Sitgreaves National Forest, August 10, 1967, L. A. Kelton, 13 (CNC). Show Low, 3 mi NNW of, July 26, 1952, A. A. Alcorn, 13 (KU). Pima Co. : Sabino Canyon, Santa Catalina Mountains , July 31, 1941, R. H. Beamer, 13 (KU). Pinal Co.: Maricopa , October 17, 1937, J. A. Kusch, 13 (CAS). Yavapai Co.: Chino Valley, July 3, 1950, R. H. Beamer, 13 (KU). Cottonwood, August 13, 1975, J. C. Schaffner, 13 (TAMU). Page Springs junction on Rt 89A, 1 mi S of, 1700 m, June 21, 1980, R. T. Schuh ( Chenopodiaceae ), 23 (AMNH). Peeples Valley, August 18, 1967, L. A. Kelton, 53 (CNC). Prescott, August 16, 1972, L. A. Kelton (CNC). Sedona, 3 mi SW of, August 18, 1968, D. E. Bright (CNC). Yuma Co. : Yuma , July 8, 1954, Deal and Swift, 13 (UCB). Arkansas: Boone Co.: 1 mi N of Omaha, August 23, 1955, A. A. Alcorn, 13 (KU). California: Fresno Co. : Coalinga, May 14, 1938, M. A. Cazier, 53 (AMNH). Fresno, July 16, 1967, T. R. Haig, 13 (CAFA). Fresno, June 3, 1956, R. O. Schuster, 13 (UCB). Inyo Co. : Alabama Hills, July 8, 1973, J. D. Pinto and G. R. Ballmer, 33 (UCR). Batcheld­ er Spring, Westguard Pass, September 7, 1969, M.

episternal scent efferent system, lateral view. E. Pretarsus, frontal view. F. Pretarsus, apical ventral view with detail of parempodia. G. Hind femur, lateral view. H. Pygophore, posterolateral view, showing left paramere and phallotheca in repose. Scales as indicated.

S. and J. Wasbauer, 13 (CAFA). Big Pine, June 16, 1925, R. L. Usinger, 463 (CAS). Big Pine, September 4, 1956, R. M. Bohart, 13 (UCD). Bishop, June 22, 1929, R. L. Usinger, 13 (CAS). Deep Springs, Deep Springs Lake, May 13, 1974, B. Villegas, 13 (UCD). Deep Springs, September 8, 1969, M. Wasbauer, 193 (CAFA). Little Lake, June 14, 1973, J. D. Pinto, 83 (UCR). Westgard Pass, 13 mi E of Big Pine, August 3, 1962, D. R. Smith, 13 (OSU). Westgard Pass, 4.5 mi W of White Mountain Road, August 22, 1963, H. B. Leech, 13 (CAS). Kern Co.: Bakersfield, July 24, 1929, R. H. Beamer, 13 (KU). Bakersfield, June 5, 1930, R. L. Usinger, 73 (UCB). Comanche Point, April 29, 1954, O. G. Bacon, 13 (UCD). Inyokern, November 7, 1969, P. H. Arnaud, Jr., 13 (CAS). Headquarters Campground, Sequoia National Forest, near Kernville, June 18, 1993, W. F. Chamberlain, 13 (TAMU). Mojave, 7 mi NE of, April 1, 1966, O’Brien, 23 (UCB). Oildale, April 3, 1983, 13 (TAMU). Poso Creek, June 5, 1929, E. P. Van Duzee, 43 (CAS). Poso Creek, June 5, 1929, R. L. Usinger, 93 (CAS). Shafter, July 9, 1940, C. G. Lewis ( Malvaceae ) (UCB). Shafter, June 12, 1931, G. L. Smith, 43 (UCB). Shafter, 13 mi S of, May 16, 1965, A. S. Menke, 13 (UCD). Tehachapi Pass, June 6, 1929, R. L. Usinger, 113 (CAS). Wofford Heights, 12 km W of, 1700 m, July 26, 1999, M. D. Schwartz, Phacelia sp. (Hydrophyllaceae) (CNC). Lassen Co.: Doyle, June 29, 1957, E. I. Schlinger, 13 (UCD). Hallelujah Junction, July 12, 1980, J. A. Skinner, 23 (UCD). Hallelujah Junction, June 29, 1976, M. A. Rabel, 23 (UCD). Hallelujah Junction, September 1, 1974, R. D. Moon, 23 (UCD). Ravendale, 10 mi S of, February 5, 1964, R. C. Dickson, 13 (UCR). Ravendale, S of, July 7, 1973, R. C. Dickson, 23 (UCR). Los Angeles Co.: Flintridge, May 20, 1955, C. L. Hogue, 13 (LACM). La Brea, June 16, 1925, L. J. Muchmore, 73 (LACM). Lancaster, 6 mi W of, November 1, 1974, J. Doyen, Chrysothamnus nauseosus (Asteraceae) (UCB). Mint Canyon, June 21, 1949, H. E. Cott, 13 (UCD). Mint Canyon, Oaks, April 20, 1932, E. P. Van Duzee, 13 (CAS). Modoc Co.: Inspection Station, 8 mi S of, Highway 139, July 17, 1955, Joe Schuh, 33 (OSU). Willow Ranch, 3 mi E of, June 6, 1970, Isatis tinctoria (Brassicaceae) (UCB). Mono Co.: Crooked Creek Naval Res. Station, White Mts. , 10,000 ft, July 7, 1966, L. and C. W. O’Brien, 13 (UCB). Sardine Creek, 8500 ft, July 8, 1951, A. T. McClay, 13 (UCD). Monterey Co.: Salinas Road at King City, May 20, 1976, Doyen, 13 (UCB). Napa Co.: Samuel Springs, May 18, 1955, E. I. Schlinger, 13 (UCD). Orange Co.: El Modena, June 6, 1935, C. E. Norland ( Asteraceae ) (LACM). Huntington, August 13, 1963, D. J. Burdick, 13 (UCB). Riverside Co.: 6 mi E of jct of highways 395 and 71, roadside, April 4, 1968, S. and S. Frommer, 13 (UCR). Aguanga, July 25, 1995, G. M. Chamberlain, 63 (TAMU). Anza, 6 mi W of, July 26, 1962, D. R. Smith, 133 (OSU). Beaumont, 5 mi E of, October 26, 1952, R. F. Smith and E. G. Linsley, Haplopappus sp. (Asteraceae) (UCB). Box Springs Mountains , July 25, 1964, G. E. Wallace, 13 (UCR). Boyd Desert Research Center, 4 mi S of Palm Desert, April 13, 1963, D. E. and A. Hardy, 13 (UCB). Cabazon, October 26, 1952, R. F. Smith and E. G. Linsley, 13 (UCB). Cathedral City, August 10, 1950, L. W. Isaak, 33 (UCD). Coyote Canyon, 3.5 mi S of Palm Desert, Boyd Desert Research Center, May 19, 1973, E. E. McGregor, 13 (UCR). Menifee Valley, hills on W end, 560 m, May 11, 1978, R. T. Schuh and J. D. Pinto, 13 (AMNH). Menifee Valley, hills on W end, 1800 ft, October 25, 1981, J. D. Pinto, 13 (UCR). Palen Dunes, April 27, 1978, Alan R. Hardy, 103 (CAFA). Palen Dunes, September 20, 1977, Hardy and F.G. Andrews, 73 (CAFA). Palm Springs, April 30, 1928, C. C. Searl, 13 (SDNH). Palm Springs, October 15, 1947, Pectis pappisa (Asteraceae) (UCR). Quail Valley, Coastal Sage Scrub Community, October 15, 1973, J. D. Pinto, 13 (UCR). Red Rock Canyon, 4 mi E of, 4 mi E of Elsinore, April 14, 1965, C. A. Tosch, 13 (UCB). Riverside, August 15, 1953, J. C. Hall, 93 (UCD). Riverside, March 27, 1963, E. I. Schlinger, 13 (UCR). Riverside, October 12, 1956, J. K. Ferguson and J. Richardson, 23 (UCR). Saboba Springs, June 2, 1917, E. P. Van Duzee, 13 (CAS). San Jacinto, April 30, 1928, C. C. Searl, 13 (SDNH). Sun City, August 28, 1968, D. E. Hardy (CNC). Thousand Palms, April 19, 1955, W. R. Richards (CNC). Whitewater Canyon, July 26, 1963, E. I. Schlinger, 13 (UCR). San Bernardino Co.: Adelanto, September 8, 1966, T. R. Haig, 13 (CAFA). Apple Valley, October 7, 1956, J. Kirk Ferguson, 13 (UCR). Cucamonga, June 8, 1945, R. C. Dickson, Franseria sp. (Asteraceae) (CAFA). Highland, 5 mi SE of, September 7, 1962, J. T. Doyen, 13 (UCB). Lucerne Valley, 4.5 mi E of, 1000 m, May 13, 1978, J. D. Pinto and R. T. Schuh, Ambrosia dumosa (Asteraceae) , 13 (AMNH). Redlands, August 16, 1955, Menke and Stage, 13 (LACM). Redlands, August 28, 1947, R. A. Flock, Chenopodium sp. (Chenopodiaceae) (UCR). Rialto, July 18, 1956, E. I. Schlinger, 13 (UCD). Yucca Valley, 10 mi N of, May 28, 1975, J. D. Pinto, Acamptopappus sphaerocephalus (Asteraceae) (UCR). San Diego Co.: Borrego Valley, June 6, 1940, R. L. Usinger, 13 (UCB). El Cajon, April 23, 1920, E. P. Van Duzee, 13 (CAS). Jacumba, August 19, 1967, L. A. Kelton (CNC). No specific locality, October 18, 1913, E. P. Van Duzee, 13 (CAS). Warner Springs, May 20, 1928, C. C. Searl, 13 (SDNH). Warner Springs, 2 mi W of, June 19, 1978, J. D. Pinto, Quercus agrifolia (Fagaceae) (UCR). San Joaquin Co.: Lodi, August 18, 1929, H. H. Keifer, 23 (CAS). Tracy, June 13, 1940, J. W. MacSwain, 13 (UCB). San Luis Obispo Co.: Arroyo Grande Creek SW of San Luis Obispo, 160 m, May 8, 1985, R. T. Schuh and B. M. Massie, Lupinus ludovicianus (Fabaceae) , 13 (AMNH). Shasta Co.: 8 mi W of Fall River Mills, 845 m, July 7, 1979, R. T. and J. Schuh, Iva axillaris (Asteraceae) , 203 (AMNH). Siskiyou Co.: Dorris, July 1, 1935, R. H. Beamer, 13 (KU). Macdoel, June 16, 1958, Joe Schuh, Iva axillaris (Asteraceae) (OSU). Trinity Co.: Junction City, June 30, 1975, T. R. Haig, 23 (CAFA). Tulare Co.: Exeter, J. K. Ellsworth, 23 (UCB). Lemon Cove, June 26, 1929, R. H. Beamer, 13 (KU). Tuolomne Co.: Strawberry, August 8, 1929, P. W. Oman, 23 (KU). Yolo Co.: Davis, August 3, 1955, A. T. McClay, 13 (UCD). Davis, July 22, 1956, A. T. McClay, 13 (UCD). Davis, April 24, 1966, R. O. Schuster, 13 (UCD). Winters, 8 mi W of, May 5, 1968, J. S. Buckett, 13 (CAFA). Woodland, June 14, 1933, E. C. Zimmerman, 13 (UCB). Colorado: Arapahoe Co.: Cherry Creek State Park, July 8, 1977, D. A. and J. T. Polhemus, 53 (AMNH). Archuleta Co.: Pagosa Springs, 7200 ft, August 24, 1972, L. A. Kelton (CNC). Pagosa Springs, 7200 ft, June 6, 1919, 13 (AMNH). Baca Co. : Springfield, June 18, 1988, R. Wharton and B. Mann, 33 (TAMU). Bent Co.: Las Animas, August 20, 1936, M. B. Jackson, 13 (KU). Boulder Co.: Boulder, August 11, 1973, G. G. E. Scudder (CNC). Boulder, June 19, 1922, 13 (AMNH). Boulder, June 21, 1903, E. P. Van Duzee, 13 (CAS). Boulder, 4 mi NW of, 6900 ft, June 8, 1961, B. H. Poole (CNC). Boulder, 6 mi W of, August 12, 1973, J. C. Schaffner, 103 (TAMU). Eldora, July 12, 1949, L. D. Beamer, 13 (KU). Nederland, Science Lodge, 9500 ft, July 4, 1961, B. H. Poole (CNC). S of St. Vrain Canyon, August 16, 1973, G. F. Knowlton and W. J. Hanson, 13 (USU). Ward, Roosevelt National Forest, July 30, 1968, L. A. Kelton, Solidago sp. (Asteraceae) (CNC). Clear Creek Co.: Doolittle Ranch, Mount Evans, 9700– 9800 ft, July 11, 1964 and August 10, 1964, B. H. Poole (CNC). Echo Lake, 10,000 ft, August 27, 1978, D. A. Polhemus, 13 (JTP). Loveland Pass, 10,000 ft, June 8, 1961, B. H. Poole (CNC). Mount Evans, 12,000 ft, August 3, 1961, B. H. Poole (CNC). Mount Evans, 11,300 ft, August 10, 1961, J. G. Chillcott (CNC). Mount Goliath, 12,000 ft, August 7, 1987, D. A. and J. T. Polhemus, 13 (JTP). Summit Lake, Mount Evans, 12800 ft, July 10, 1961, W. R. M. Mason (CNC). Timberline, Mount Evans, 11,600 ft, July 13, 1961, J. R. Stainer (CNC). West Chicago Creek, Arapaho National Forest, 9800 ft, July 28, 1968, L. A. Kelton (CNC). Custer Co.: 7 mi W of Greenwood, August 11, 1964, J. G., B. L., and K. C. Rozen, 13 (AMNH). Douglas Co.: Chatfield State Park, June 18, 1978, J. T. Polhemus, 13 (JTP). Daniels Park, July 2, 1979, D. A. Polhemus, 13 (JTP). Perry Park, July 13, 1977, D. A. and J. T. Polhemus, 23 (JTP). Roxborough Road near Chatfield Park, July 9, 1978, J. T. Polhemus, 13 (JTP). S of Roxborough Park, August 24, 1982, D. A. and J. T. Polhemus, 23 (JTP). Waterton, August 8, 1987, T. J. Henry, Artemisia sp. (Asteraceae) , 53, 1♀ (USNM). Eagle Co.: Gypsum, 6300 ft, August 28, 1959, E. Ordway, 13 (KU). El Paso Co.: Calhan, August 19, 1936, R. H. Beamer, 13 (KU). Colorado Springs, 5915 ft, E. S. Tucker, 63 (KU). Colorado Springs, June 27, 1966, J. A. Slater, 13 (AMNH). Security, July 4, 1983, J. T. Polhemus, 13 (JTP). Elbert Co.: 5 mi E of Agate, August 9, 1987, J. T. and D. A. Polhemus, 13 (JTP). Garfield Co.: Glenwood Springs, 5800 ft, July 22, 1920 and August 5, 1920, 63 (AMNH). Gunnison Co.: Gunnison, August 30, 1954, A. A. Alcorn, 13 (KU). Huerfano Co.: 3 mi W of Walensburg, near stream, 6500 ft, August 17, 1982, L. Herman, 93 (AMNH). Jackson Co.: Rabbit Ears Pass, June 3, 1947, R. H. Beamer, 13 (KU). Jefferson Co.: Deer Creek Canyon, 6500 ft, July 11, 1986, R. T. Schuh and J. T. Polhemus, Quercus gambelii (Fagaceae) (AMNH). Morrison, 5766 ft, July 27, 1968, L. A. Kelton (CNC). Waterton, Platte River, 5350 ft, July 11, 1986, R. T. Schuh and J. T. Polhemus, 13 (AMNH). La Plata Co.: Durango, August 26, 1972, L. A. Kelton, Chenopodium album (Chenopodiaceae) (CNC). Larimer Co.: Chambers Lake, Roosevelt National Forest, 9200 ft, August 11, 1968, L. A. Kelton (CNC). Fall River Pass, Rocky Mountain National Park, August 17, 1936, R. H. Beamer, 13 (KU). Glen Haven, July 12, 1946, P. B. Lawson, 13 (KU). Hudson, August 25, 1925, C. J. Drake, 13 (USNM). Owl Canyon, 6000 ft, August 15, 1968, L. A. Kelton (CNC). Rocky Mountain National Park, August 5, 1947, L. D. Beamer, 13 (KU). Las Animas Co.: No specific locality, September 22, 1927, P. A. Readio, 13 (KU). Mesa Co. : Fruita, 4500 ft, July 16, 1919, 363 (AMNH). Grand Junction, September 4, 1938, D. E. Hardy, 23 (KU). Moffat Co.: Massadona, August 8, 1965, G. F. Knowlton, 33 (USU). Montezuma Co. : Cortez, July 19, 1968, L. A. Kelton, Salicornia sp. (Chenopodiaceae) (CNC). Montrose Co.: 15 mi SW of Montrose, August 8, 1987, T. J. Henry, 13 (USNM). Otero Co.: La Junta, May 13, 1979, J. T. and D. A. Polhemus, 13 (JTP). Pitkin Co.: Aspen, 8000 ft, July 24, 1919, 13 (AMNH). Pueblo Co.: Pueblo, 5000 ft, August 19, 1960, R. E. Dreisbach, 13 (USNM). Rio Blanco Co. : Meeker, 6200 ft, July 20, 1919, 13 (AMNH). Routt Co.: Steamboat Springs, 6900 ft, July 11, 1964, H. H. Knight (CNC). Steamboat Springs, July 9, 1964, H. H. Knight, 13 (USNM). San Juan Co.: San Juan National Forest, Chimney Rock, July 15, 1991, W. F. Chamberlain, 13 (TAMU). Unknown Co.: Little Beaver Creek, July 11, 1937, C. L. Johnston, 73 (KU). Macedonia, July 1, 1931, H. T. Peters, 23 (KU). Regnier, 4500 ft, June 6, 1919, 13 (AMNH). Weld Co.: Keenesburg, June 11, 1961, B. H. Poole (CNC). Pawnee National Grassland Headquarters, August 9, 1973, G. G. E. Scudder (CNC). Nunn, 17.5 mi N of, Pawnee National Grassland, August 6, 1973, J. A. Slater and R. Baranowski, 13 (AMNH). Rockport, 3 mi N of, 5794 ft, July 23, 1971, O’Brien and Marshall, 13 (AMNH). Roggen, July 2, 1978, G. E. Bohart, 13 (USU). Yuma Co. : 13.5 mi E of Idalia, August 10, 1964, J. G. and B. L. Rozen, 13 (AMNH). Wray, 3700 ft, August 17, 1919, 23 (AMNH). Connecticut: New Haven Co.: Mount Carmel, August 8, 1962, A. Decaprio, 13 (AMNH). New Haven, August 20, 1934, R. H. Beamer and M. E. Griffith, 143 (KU). N of Southbury on Rt 67, September 1, 1970, F. P. Maroney, 13 (AMNH). Tolland Co.: Somers, LaChance Farm, July 8, 1965, J. A. Slater, 13 (AMNH). Storrs, July 3, 1954, J. A. Slater, 63 (AMNH). Florida: Seminole Co.: No specific locality, June 10, 1900, E. M. Davis, 13 (AMNH). Idaho: Bear Lake Co.: Bloomington Lake, T14S, R42E, sec. 5, July 24, 1982, M. D. Schwartz, Picea engelmannii (Pinaceae) (AMNH). Blaine Co.: 20 mi N of Shoshone, Big Wood River, July 1, 1968, W. Gagne, J. Haddock, 93 (UCB). Bonneville Co.: Swan Valley, 4 mi W, 5400 ft, July 10, 1973, P. Oman, 13 (OSU). Cassia Co. : Burley, May 6, 1931, R. H. Beamer, H. T. Peters, M. W. Sanderson, 63 (KU). Franklin Co.: Cub River Canyon, August 28, 1954, G. F. Knowlton, 13 (KU). Latah Co.: 5 mi E of Harvard, Paillasse River, July 3, 1966, W. Gagne, J. Haddock (CNC). New Perce Co.: Lewiston, August 3, 1972, L. A. Kelton, Medicago sativa (Fabaceae) (CNC). Oneida Co. : Holbrook Summit, August 20, 1969, G. F. Knowlton, 23 (USU). Owyhee Co.: 40 mi S of Bruneau, Clover Creek, June 20, 1982, T. J. Henry, Iva axillaris (Asteraceae) , 83, 4♀ (USNM). Power Co.: American Falls, August 5, 1972, L. A. Kelton (CNC). Illinois: Boone Co.: Belvedere, August 7, 1950, J. A. Slater, 13 (AMNH). Champaign Co.: Champaign, August 9, 1937, Mohr and Burke, 13 (AMNH). Champaign, October 3, 1956, J. F. McAlpine (CNC). Urbana, October 7, 1941, J. A. Slater, 13 (INHS). Hancock Co.: Hamilton, July 16, 1967, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Nauvoo, July 17, 1967, L. A. Kelton (CNC). McLean Co.: Bloomington, July 24, 1971, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Mercer Co.: Norwood, July 16, 1967, L. A. Kelton (CNC). Sangamon Co.: Springfield, September 16, 1932, T. H. Frison, 13 (AMNH). Tazewell Co.: Delavan, July 12, 1932, Dozier and Park, 13 (AMNH). Warren Co.: Roseville, August 20, 1940, R. I. Sailer, 13 (KU). Winnebago Co.: Rockford, September 2, 1930, T. H. Frison, 13 (AMNH). Indiana: Howard Co.: NW Howard County, August 20, 1984, D. A. Rider, 13 (DAR). Kosciusko Co.: 2 mi N of Warsaw, September 1, 1966, C. W. O’Brien, 13 (UCB). Iowa: Adair Co.: Adair, July 25, 1960, P. M. Marsh, 13 (UCD). Boone Co.: Boone, May 22, 1949, J. A. Slater, 13 (AMNH). Ledges State Park, July 9, 1956, J. C. Schaffner, 13 (TAMU). Clay Co.: Peterson, June 13, 1981, D. A. Polhemus, 13 (JTP). Dickinson Co.: Silver Lake, July 8, 1963, J. C. Schaffner, 13 (TAMU). Henry Co.: 7 mi W of Mount Pleasant, July 2, 1976, J. C. Schaffner, 13 (TAMU). Mahaska Co.: New Sharon, July 17, 1932, F. Andre, 13 (UCB). Polk Co.: 4 mi NE of Alleman, Polk City, May 28, 1950, J. Laffoon, 13 (AMNH). Story Co.: Ames, August 16, 1934, H. A. Scullen, 13 (OSU). Ames, August 23, 1952, J. C. Schaffner, 63 (TAMU). Ames, August 9, 1932, F. Andre, 13 (UCB). Ames, Exp. Station, June 4, 1897, E. D. Ball, 13 (UAZ). Ames, June 1, 1954, L. A. Kelton (CNC). Ames, June 13, 1950, J. A. Slater, 23 (AMNH). Ames, October 1, 1949, W. Downes, 13 (OSU). 4 mi E of Gilbert, May 11, 1949, J. Laffoon, 13 (AMNH). Pleat Bog Valley, Ames, July 22, 1928, G. S. Walley (CNC). Sopers Mill Dam, 4 mi E of Gilbert, May 24, 1949, J. A. Slater, 23 (AMNH). Warren Co.: 1.5 mi E of Hartford, July 5, 1976, J. C. Schaffner, 13 (TAMU). Kansas: Crawford Co.: Cherokee, September 2, 1941, R. H. Beamer, 13 (KU). Douglas Co.: Lawrence, August 10, 1936, L. J. Lipovsky, 23 (KU). Lawrence, August 14, 1954, W. E. LaBerge, 13 (KU). Lawrence, October 12, 1963, B. Harrison, 13 (KU). No specific locality, August 22, 1920, W. K. Hoffmann, 23 (KU). Finney Co.: Garden City, June 12, 1940, R. H. Beamer, 13 (KU). Garden City, Sand H., September 10, 1935, M. E. Griffith, 103 (KU). Garden City, September 3, 1951, C. D. M. and W. E. L., 13 (KU). Kearny Co.: No specific locality, 2998 ft, July 15, 1921, 213 (KU). Kingman Co.: No specific locality, 1516 ft, R. H. Beamer, 13 (KU). Meade Co.: No specific locality, September 13, 1944, R. H. Beamer, 13 (KU). Osborne Co.: No specific locality, 13 (KU). Pratt Co.: Cairo, June 11, 1964, Kerfoot and Michener, Gaillardia pulchella (Asteraceae) (KU). No specific locality, 1900 ft, June 12, 1921, 13 (KU). Rawlins Co.: No specific locality, 2850 ft, F. X. Williams, 13 (KU). Reno Co.: Hutchinson, September 7, 1938, D. E. and A. Hardy, 13 (KU). Medora, sand dunes, June 13, 1934, R. H. Beamer, 13 (KU). No specific locality, September 3, 1919, W. E. Hoffmann, 463 (KU). Republic Co.: No specific locality, July 11, 1935, R. H. Beamer, 13 (KU). Scott Co.: Lake Scott State Park, July 26, 1992, E. G. Riley, 13 (TAMU). Sedgwick Co.: Wichita, June 21, 1940, D. E. Hardy, 33 (KU). Seward Co.: Liberal, August 16, 1945, R. H. Beamer, 13 (KU). Liberal, June 22, 1940, D. E. Hardy, 23 (KU). No specific locality, October 28, 1945, R. H. Beamer ( Malvaceae ) (KU). Sheridan Co.: Hoxie (near), August 21, 1950, V. Roth, 23 (OSU). Smith Co.: No specific locality, July 8, 1926, R. H. Beamer, 13 (KU). Massachusetts: Essex Co.: Beach Bluff, July 24, 1914, H. M. Parshley, 123 (CAS). Swampscott, August 19, 1914, H. M. Parshley, 13 (CAS). Middlesex Co.: Holliston, August 1, 1900, N. Banks, 103 (AMNH). Norfolk Co.: Needham, September 3, 1911, C. W. Johnson, 13 (CAS). Wellesley, July 11, 1909, E. P. Van Duzee, 13 (CAS). Suffolk Co.: Forest Hills, September 19, 1914, H. M. Parshley, 13 (CAS). Michigan: Cheboygan Co.: No specific locality, July 27, 1933, H. B. Hungerford, 13 (KU). Emmet Co.: 6 mi S of Pellston, July 27, 1965, W. R. Anderson, 13 (AMNH). Monroe Co.: Temperance, September 10, 1963, G. G. E. Scudder (CNC). Shiawassee Co.: T5N, R1E, sec. 20–29, May 28, 1967, R. T. Schuh, T. Hlavac, 13 (AMNH). Minnesota: Chisago Co.: North Branch, August 5, 1922, H. H. Knight, 103 (USNM). Clay Co.: Bluestem SNA D6, T139N, R46W, sec. 15, August 22, 1995, Rider, Locken, Fauske, 63 (AMNH). Polk Co.: Agassiz Dunes, T147N, R44W, sec. 32, August 28, 1995, Marty, Urlacher, Locken, 203 (DAR). Crookston, July 11, 1953, B. A. Haws, 43 (USU). Ramsey Co.: Saint Anthony Park, July 25, 1921, H. H. Knight, 23 (USNM). Saint Paul, University Farm, August 5, 1921, H. H. Knight, 53 (USNM). Renville Co.: Olivia, June 28, 1921, H. H. Knight, 13 (USNM). Scott Co.: dunes near Jordan, August 1, 1923, W. H. Hoffmann, 13 (USNM). Mississippi: Unknown Co.: Luka, July 14, 1930, P. W. Oman, 13 (KU). Missouri: Clinton Co.: Lathrop, June 27, 1955, E. C. Becker (CNC). Jackson Co.: Atherton, July 10, 1915, C. F. Adams, 23 (CAS). Atherton, May 19, 1922, C. F. Adams, 13 (KU). Kansas City, F. Rogers, 53 (KU). Randolph Co.: 1 mi E of Moberly, July 4, 1986, E. G. Riley, 13 (DAR). Montana: Gallatin Co.: Elkhorn Ranch, Gallatin River, Upper Gallatin Canyon, 7000 ft, August 1, 1928, J. McDunnough (CNC). Glacier Co.: East Glacier Park, 6 mi N on Rt 49, Looking Glass Hill, August 3, 1994, M. D. Schwartz, 13 (AMNH). Jefferson Co.: Whitehall, August 13, 1931, H. T. Peters, 23 (KU). Mineral Co.: St. Regis , 770 m, August 28, 1981, P. H. Arnaud, Jr., 13 (CAS). Nebraska: Adams Co.: Hastings, June 1, 1900, A. P. Morse, 23 (AMNH). Antelope Co. : Royal, July 27, 1973, L. A. Kelton (CNC). Brown Co.: extreme NW corner of Brown County, June 10, 1950, Hicks, Slater, and Laffoon, 13 (AMNH). Cass Co.: rest area on I­ 80, 2 mi W of Platte Rd., June 29, 1980, K. Schmidt, 13 (AMNH). Dawson Co.: Cozad, July 23, 1960, P. M. Marsh, 13 (UCD). Douglas Co.: Omaha, May 14, 1936, K. Rankin, 13 (KU). Hall Co.: Grand Island, July 8, 1964, H. H. Knight, 23 (CNC, USNM). Hitchcock Co.: 4 mi W of Culbertson, August 10, 1964, J. G. and B. L. Rozen, 13 (AMNH). Keith Co.: Lake McConaughy, July 21, 1973, W. F. Chamberlain, 13 (TAMU). Platt Co. : No specific locality, July 28, 1949, D. G. Denning, 13 (CNC). Red Willow Co.: Indianola, July 5, 1900, A. P. Morse, 53 (AMNH). Sioux Co.: 7 mi N of Harrison, August 13, 1962, J. G. and B. L. Rozen, 43 (AMNH). 8 mi N of Harrison, August 9, 1971, J. G., B. L., and K. C. Rozen, 73 (AMNH). Nevada: Carson City Co. : Carson City, June 27, 1929, R. L. Usinger, 13 (CAS). Churchill Co.: Fallon, August 12, 1940, D. E. Hardy, 23 (KU). Elko Co.: Carlin, June 25, 1935, R. H. Beamer, 13 (KU). Humboldt Co.: Orovada, June 15, 1960, A. S. Menke, 13 (UCD). Lander Co.: Austin, August 12, 1940, D. E. Hardy and R. H. Beamer, 63 (KU). Lyon Co.: Fernley, August 20, 1980, J. Powell, J. DeBenedictis, 13 (UCB). Pershing Co.: 7 mi E of Oreana, August 12, 1976, T. R. Haig, 13 (CAFA). Washoe Co.: Nixon, August 14, 1963, R. L. Westcott, 23 (LACM). Patrick, June 30, 1964, J. A. Froebe, 13 (UCD). Pyramid Lake, June 27, 1972, R. A. Belmont, 23 (UCD). Reno, July 3, 1929, F. E. Blaisdell, 13 (CAS). Wadsworth, August 30, 1967, L. A. Kelton, Ambrosia sp. (Asteraceae) (CNC). Wadsworth, 2 mi W of, June 23, 1962, G. I. Stage, 13 (UCB). White Pine Co. : Charcoal Ovens State Park, July 5, 1960, T. R. Haig, 33 (UCD). Ely, August 13, 1940, R. H. Beamer, 23 (KU). New Hampshire: Coos Co.: Lake of the Clouds, Mount Washington, 500 ft, August 1, 1954, Becker, Munroe, Mason (CNC). Mount Washington, A. T. Slosson, 13 (AMNH). Grafton Co.: Franconia, A. T. Slosson, 13 (AMNH). New Jersey: Bergen Co.: Ridgewood, August 30, 1969, 13 (AMNH). Passaic Co.: Great Notch, August 20, 1925, Scott, 13 (AMNH). New Mexico: Bernalillo Co.: Albuquerque, June 5, 1941, R. H. Beamer, 63 (KU). Catron Co.: 18 mi E of Quemado, August 15, 1982, J. C. Schaffner, 153 (TAMU). Chaves Co.: Elkins, July 20, 1967, L. A. Kelton (CNC). Colfax Co. : 5 mi E of Eagle’s Nest, August 8, 1969, J. C. Schaffner, 13 (TAMU). Eddy Co.: No specific locality, May 30, 1979, Burke, Schaffner, et al., 33 (TAMU). Grant Co. : Silver City, 1800 m, August 1, 1977, N. L. H. Krauss, 13 (AMNH). Silver City, 1800–1900 m, August 15, 1977, N. L. H. Krauss, 33 (AMNH). Guadalupe Co.: Cuervo, June 23, 1940, L. J. Lipovsky, 73 (KU). Santa Rosa , June 23, 1940, D. E. Hardy, 23 (KU). Santa Rosa , June 23, 1941, L. H. Banker, 13 (KU). Hidalgo Co.: 18 mi N of Rodeo, August 25, 1958, G. B. Pittman, 13 (UCD). Lea Co. : No specific locality, September 24, 1979, R. R. Murray and J. C. Schaffner, 43 (TAMU). Otero Co.: Cloudcroft, June 27, 1940, L. J. Lipovsky, 53 (KU). Mayhill, July 23, 1967, L. A. Kelton (CNC). Cloudcroft, 2 mi S of, July 17, 1979, Delorme, McHugh, Schaffner, 13 (TAMU). Cloudcroft, 2.5 mi E of, June 28, 1973, W. N. Mathis, 13 (OSU). Cloudcroft, 4 mi E of, September 26, 1979, Murray and Schaffner, 13 (TAMU). Cloudcroft, 7 mi N of, August 13, 1982, J. C. Schaffner, 33 (TAMU). Mountain Park, July 27, 1940, L. J. Lipovsky, 13 (KU). Tularosa, 25 mi W of, July 1, 1940, D. E. Hardy, 23 (KU). Roosevelt Co.: Oasis State Park, 4 mi N, 2 mi W of Portales, stabilized dune, July 12, 1977, S. Frommer, 13 (UCR). San Miguel Co.: Conchas Dam, 15 mi SW of, July 16, 1991, G. M. Chamberlain, 33 (TAMU). Ilfeld, July 3, 1968, L. A. Kelton, Helianthus sp. (Asteraceae) (CNC). Montezuma, Gallinas Canyon , August 22, 1972, L. A. Kelton (CNC). Trujillo, July 3, 1968, L. A. Kelton (CNC). Socorro Co. : Magdalena, Cibola National Forest, August 21, 1972, L. A. Kelton (CNC). Magdalena, 12 mi W of, August 15, 1982, J. C. Schaffner, 13 (TAMU). No specific locality, August 18, 1927, R. H. Beamer, 43 (KU). Torrance Co.: Estancia, June 24, 1940, D. E. Hardy, 53 (KU). Manzano, June 26, 1941, B. Hogden, 23 (KU). Manzano, 6 mi SW of Red Canyon Campground, June 11, 1982, J. D. Pinto, 13 (UCR). Tajique, June 25, 1940, R. H. Beamer, 23 (KU). New York: Columbia Co. : Chatham, August 23, 1904, A. P. Morse, 53 (AMNH). Erie Co.: Buffalo, August 24, 1898, E. P. Van Duzee, 33 (CAS). Buffalo, June 9, 1888, E. P. Van Duzee, 13 (AMNH). Essex Co.: Whiteface Mountain, 4600–4872 ft, July 19, 1962, J. R. Vockeroth (CNC). Genesee Co.: Batavia, July 31, 1915, H. H. Knight, 33, 1♀ (CAS, CNC). Batavia, September 2, 1915, H. H. Knight, 13 (USNM). Nassau Co.: Sea Cliff, Long Island, N. Banks, 23 (AMNH). Rockland Co.: Pearl River, June 21, 1914, 13 (AMNH). Suffern, September 17, 1910, 13 (AMNH). Suffolk Co.: Huntington, Long Island, Kalbfleisch Research Station, August 16, 1962, P. H. Arnaud, 13 (AMNH). Tompkins Co.: Ithaca, September 4, 1973, M. D. Schwartz, 13 (AMNH). Warren Co.: Warrensburg, August 25, 1955, R. Baranowski, 33 (AMNH). North Dakota: Bottineau Co.: 10 mi S of Antler, Walker Farm, July 30, 1920, A. C. Burrill, 23 (AMNH). Cass Co.: Fargo, July 31, 1977, N. L. H. Krauss, 63 (AMNH). Grand Forks Co.: 6 mi NW of Grand Forks, August 19, 1993, M. D. Schwartz, Helianthus sp. (Asteraceae) (CNC). Richland Co.: Sheyenne National Grassland, 2.5 mi SE of McLeod, May 26, 1992, D. A. Rider, 13 (DAR). Ohio: Summit Co.: Barberton, August 10, 1936, L. J. Lipovsky, 143 (KU). No specific locality, August 3, 1936, L. J. Lipovsky, 13 (KU). Unknown Co.: Trading Post, September 10, 1963, G. G. E. Scudder (CNC). Oregon: Baker Co.: 3 mi E of Baker, Conrad Allen Ranch, August 6, 1963, C. W. Baker, 23 (OSU). Hells Canyon, Ballards Landing, July 31, 1976, J. D. Lattin, 23 (OSU). Crook Co.: 15 mi SE of Prineville, near Eagle Rock, June 9, 1968, P. Oman, 103 (OSU). Douglas Co.: Roseburg, 71 mi E, 3400 ft, July 26, 1945, H. A. Scullen, 103 (OSU). Grant Co. : Izee, Keerin’s Ranch, June 25, 1935, Joe Schuh, 13 (AMNH). Izee, Keerin’s Ranch, June 25, 1955, L. A. Kelton (CNC). Seneca, June 30, 1955, Joe Schuh, 13 (AMNH). Harney Co.: Fish Lake, Steens Mts., 7200 ft, August 12, 1958, K. M. Fender, 13 (OSU). Frenchglen, 1 mi E of, July 10, 1968, J. D. Lattin, 13 (OSU). Malheur Wildlife Refuge, 10 mi S of headquarters, July 12, 1968, Oman, 53 (OSU). T36S, R35E, sec. 8 NW, August 8, 1969, T. Seibert and N. Cobb, 13 (OSU). Jackson Co.: Medford, 10 mi NE of, October 7, 1965, K. Goeden, 13 (OSU). Medford, 12 mi NW of, August 7, 1966, K. Goeden ( Asteraceae ) (OSDA, OSU). Pinehurst, 3400 ft, June 24, 1971, P. Oman, 13 (OSU). Siskiyou, June 14, 1959, L. A. Kelton, R. Madge (CNC). Jefferson Co.: 10 mi N of Warm Springs, July 2, 1968, Oman, 73 (OSU). Klamath Co.: between Worden and Keno, 1280 m, June 27, 1979, R. T. and J. Schuh, 13 (AMNH). Klamath Falls, July 3, 1953, J. Schuh, Trifolium sp. (Fabaceae) (CNC). Midland, August 7, 1955, Joe Schuh, Iva axillaris (Asteraceae) (OSU). Skookum Meadow, R9E, T27S, sec. 16, July 17, 1979, Oman, 13 (OSU). Lake Co.: Hart Mountain, 6250 ft, September 2, 1977, Joe Schuh ( Asteraceae ), 13 (AMNH). Lakeview, June 27, 1951, B. Malkin, 13 (CAS). Malheur Co.: Ontario, July 23, 1963, S. Besse, 13 (OSU). Morrow Co.: Boardman, May 11, 1960, K. Goeden, 33 (OSU). Boardman, May 24, 1938, K. Gray and J. Schuh, 13 (AMNH). Boardman, September 13, 1949, V. Roth, 13 (OSU). Umatilla Co.: Cayuse, July 13, 1938, Schuh and Gray, 13 (AMNH). Echo, May 11, 1960, K. Goeden ( Fabaceae ) (OSU). Wasco Co.: 5 mi W of The Dalles, May 18, 1972, J. D. Lattin, 13 (OSU). Pennsylvania: Monroe Co.: Delaware Water Gap, A. T. Slosson, 23 (AMNH). Washington Co.: Taylorstown exit off Hwy 70, July 3, 1966, P. H. Arnaud, Jr., 13 (CAS). Westmoreland Co.: Greensberg, 23 (CAS). South Dakota: Custer Co.: Custer State Park, Black Hills, August 7, 1971, L. A. Kelton (CNC). Custer, August 26, 1935, N. W. Sanderson, 13 (KU). Flynn Creek, 8 mi N of Pringle, 5400 ft, July 8, 1961, H. and A. Howden (CNC). Davison Co.: Mitchell, September 12, 1963, G. G. E. Scudder (CNC). Jones Co. : Capa, June 1, 1921, H. C. Severin, 13 (CAS). Stanley Co.: Fort Pierre, September 21, 1920, H. C. Severin, 23 (CAS). Pierre, July 19, 1937, C. L. Johnston, 13 (KU). Union Co.: 3 mi S of Jefferson, June 11, 1949, Slater and Laffoon, 23 (AMNH). Texas: Bailey Co.: 5 mi SW of Muleshoe, June 17, 1992, W. F. Chamberlain, 33 (TAMU). Carson Co.: Panhandle, July 2, 1968, L. A. Kelton (CNC). Culberson Co.: 14 mi N of Van Horn, August 11, 1982, J. C. Schaffner, 13 (TAMU). 14 mi N of Van Horn, August 11, 1982, J. C. Schaffner, Parthenium incanum (Asteraceae) , 283 (TAMU). Eastland Co.: No specific locality, August 18, 1920, Grace Olive Wiley, 23 (KU). El Paso Co.: Clint, August 16, 1976, W. F. Chamberlain, 33 (TAMU). Hartley Co.: Romero, June 22, 1940, L. J. Lipovsky, 33 (KU). Kerr Co.: Kerrville, RNC, June 15, 1996, W. F. Chamberlain, 13 (TAMU). Oldham Co.: Adrian, September 6, 1951, A. T. McClay, 33 (UCD). Parm­ er Co.: 8 mi SW of Friona, July 30, 1937, Kansas University Mex. Expedition, 13 (KU). Rusk Co.: Big Springs, June 20, 1947, A. C. Michener, 33 (KU). Ward Co.: 1 mi S of Grandfalls, April 19, 1985, J. C. Schaffner, 13 (TAMU). Utah: Box Elder Co. : Bear River City, August 16, 1953, G. F. Knowlton ( Apiaceae ) (USU). Corinne, August 8, 1961, G. F. Knowlton, 13 (KU). Corinne, July 25, 1953, G. F. Knowlton, 13 (USU). Corinne, September 22, 1949, G. F. Knowlton, 23 (USU). Deweyville, June 30, 1967, G. F. Knowlton, 23 (UCD). Deweyville, September 15, 1953, G. F. Knowlton, 23 (USU). Garland, August 25, 1953, G. F. Knowlton, 43 (USU). Garland, September 6, 1925, G. F. Knowlton, 13 (USU). Hansel Mountains, July 8, 1969, G. F. Knowlton, 83 (USU). Hardup, June 18, 1964, G. F. Knowlton, 13 (USU). Honeyville, September 16, 1948, G. F. Knowlton, Apium graveolens (Apiaceae) (USU). Kelton, June 30, 1969, G. F. Knowlton, 13 (USU). Kelton, 9 mi NW of, June 30, 1969, G. F. Knowlton and W. J. Hanson, 23 (USU). Penrose, August 20, 1949, G. F. Knowlton, 13 (USU). Snowville, August 12, 1969, G. F. Knowlton, 33 (USU). Snowville, 6 mi W of, June 27, 1969, G. F. Knowlton, 13 (USU). Tremonton, August 8, 1953, G. F. Knowlton ( Solanaceae ) (USU). Cache Co.: Avon, August 16, 1967, G. F. Knowlton, 13 (UCD). Clarkston, July 1, 1946, G. F. Knowlton ( Solanaceae ) (USU). Lewiston, August 26, 1925, G. F. Knowlton ( Asteraceae ) (USU). Logan Canyon, September 12, 1973, G. F. Knowlton, 13 (USU). Logan, August 14, 1953, W. J. Hanson and G. F. Knowlton ( Solanaceae ) (USU). Logan, July 23, 1963, G. F. Knowlton, 13 (USU). Logan, September 20, 1939, G. F. Knowlton, 13 (USU). Petersburg, September 5, 1958, K. C. Tilley, 53 (USU). Richmond, July 11, 1958, K. C. Tilley ( Fabaceae ) (USU). Wellsville, August 15, 1958, K. C. Tilley ( Fabaceae ) (USU). Davis Co.: Farmington, July 9, 1953, G. F. Knowlton and W. J. Hanson ( Apiaceae ) (USU). Farmington, June 22, 1932, G. F. Knowlton and M. J. Janes, 13 (USU). Kings Station, July 24, 1922, H. H. Knight, 33 (CAS). North Farmington, September 19, 1949, G. F. Knowlton and D. McComb ( Apiaceae ) (USU). Duchesne Co.: Duchesne, August 25, 1966, L. and C. W. O’Brien, 13 (UCB). Myton, July 15, 1953, Knowlton and Liebmann, 13 (USU). Myton, July 21, 1954, G. F. Knowlton, 33 (KU). Roosevelt, July 20, 1954, G. F. Knowlton, 23 (AMNH). Roosevelt, June 12, 1963, G. F. Knowlton, 13 (USU). Emery Co.: Hanksville ‘‘dunes’’, 17 mi N near Butte well, July 26, 1978, Andrews and Hardy, 13 (CAFA). Woodside, September 4, 1938, D. E. and A. Hardy, 13 (KU). Garfield Co.: 14.3 mi S of Rt 95 on Rt 276, 3.4 mi N of Starr Springs Turnoff, 5000 ft, June 19, 1983, R. T. Schuh and M. D. Schwartz, 103 (AMNH). Grand Co.: Arches National Park, September 3, 1947, G. F. Knowlton, 13 (USU). Green River, July 14, 1954, G. F. Knowlton, 43 (AMNH). Kane Springs, 15 mi SE of Moab, August 9, 1957, C. D. Michener, Cleome lutea (Capparidaceae) (KU). Moab, August 9, 1963, G. F. Knowlton, 13 (USU). Moab, July 14, 1991, W. F. Chamberlain, 13 (TAMU). Moab, July 15, 1954, G. F. Knowlton, 343 (AMNH). South Beaver Mesa, La Sal Mts. , July 4, 1980, J. T. and D. A. Polhemus, 13 (JTP). Iron Co. : 22 mi W of St. George, 4500 ft, March 20, 1971, Oman, 13 (OSU). Juab Co.: Levan, August 20, 1963, G. F. Knowlton, 13 (UCD). Mona Lake, July 28, 1957, A. H. Barnum, 23 (TAMU). Kane Co.: Kanab, August 11, 1965, R. S. Roberts, 13 (USU). Kanab, August 9, 1936, R. H. Beamer, 143 (KU). Millard Co.: Abraham, July 23, 1965, G. F. Knowlton, 13 (USU). Cove Fort, September 8, 1954, G. F. Knowlton, 33 (KU). Fillmore, July 22, 1965, G. F. Knowlton, 13 (USU). No specific locality, September 14, 1965, G. F. Knowlton, 13 (USU). Oak City, July 23, 1965, G. F. Knowlton, 13 (USU). Price, July 13, 1954, G. F. Knowlton, 33 (AMNH). Piute Co.: Circleville, August 11, 1957, C. D. Michener, Cleome serrulata (Capparidaceae) (KU). Price Co.: No specific locality, August 10, 1957, C. D. Michener, 13 (KU). Salt Lake Co.: Emigration Canyon, June 17, 1933, O. H. Swezey, 13 (CAS). Hunter, July 9, 1954, G. F. Knowlton, 43 (AMNH). Salt Lake City, July 20, 1948, G. F. Knowlton ( Apiaceae ) (USU). Salt Lake City, June 10, 1952, G. F. Knowlton ( Solanaceae ) (USU). Sandy, May 27, 1930, G. F. Knowlton, 13 (USU). San Juan Co.: La Sal Mts., nr Brumley Creek, T27S, R24E, 8000 ft, June 11, 1982, M. D. Schwartz, Artemisia tridentata (Asteraceae) , 13 (AMNH). Mexican Hat, July 13, 1967, G. F. Knowlton, 43 (UCD). Mexican Hat, 7.7 mi N of, on Rt 261, T41S, R18E, 5000 ft, June 17, 1983, R. T. Schuh and M. D. Schwartz, 13 (AMNH). Monticello, July 15, 1952, G. E. Bohart and G. F. Knowlton, 13 (USU). Sevier Co.: Richfield, July 28, 1954, G. F. Knowlton, 13 (AMNH). Salina, July 2, 1943, G. F. Knowlton, 33 (KU). Salina, June 24, 1954, G. L. Nielson, 63 (USU). Venice, August 17, 1961, G. F. Knowlton, Silphium sp. (Asteraceae) (KU). Summit Co.: Kimball Junction, August 31, 1967, L. A. Kelton, Chrysothamnus sp. (Asteraceae) (CNC). Tooele Co.: Erda, August 18, 1953, G. F. Knowlton, 13 (USU). Tooele, August 8, 1952, G. F. Knowlton, 13 (USU). Uintah Co.: Bonanza, 5–10 mi SW of, T11S, R24E, sec. 6, 5000–5600 ft, June 9, 1981, M. D. Schwartz, Artemisia tridentata (Asteraceae) , 13 (AMNH). Escalate Ranch, Vernal, August 21, 1954, J. B. Karren, 13 (USU). Jensen, 10 mi E of, July 13, 1993, W. F. Chamberlain, Helianthus sp. (Asteraceae) , 43 (TAMU). Vernal, 7 mi N of, August 25, 1966, L. and C. W. O’Brien, 13 (UCB). Unknown Co.: Antelope , July 1, 1931, J. Nottingham, 23 (KU). Utah Co.: American Fork, August 28, 1958, G. F. Knowlton ( Apiaceae ) (USU). American Fork, July 12, 1949, G. F. Knowlton ( Apiaceae ) (USU). Lehi, August 18, 1953, G. F. Knowlton ( Apiaceae ) (USU). Lehi, July 25, 1927, G. F. Knowlton, 13 (USU). Pleasant Grove, June 22, 1953, G. F. Knowlton, 43 (USU). Provo Canyon, August 7, 1954, G. L. Nielson, 113 (USU). Provo, environs, September 9, 1974, L. J. Westover, 13 (TAMU). Spanish Fork Canyon, July 13, 1954, G. T. Knowlton and D. W. Davis, 23 (KU). Spanish Fork, August 14, 1940, D. E. Hardy, 13 (KU). Springville, June 5, 1930, G. F. Knowlton, 13 (USU). Springville, September 23, 1953, G. F. Knowlton, 23 (USU). Thistle, July 6, 1954, G. F. Knowlton, 53 (KU). Thistle, August 12, 1947, G. F. Knowlton, 13 (USU). Washington Co.: Dixie State Park, July 28, 1965, M. S. Lee, 13 (USU). Pine Valley, June 14, 1931, B. A. Haws, 13 (USU). Pintura, August 11, 1929, R. H. Beamer, 23 (KU). St. George, 2800 ft, September 9, 1954, G. F. Knowlton, 13 (KU). Toquerville, June 18, 1935, G. F. Knowlton, 13 (USU). Zion National Park at Lava Point Campground, 7000 ft, May 24, 1982, M. D. Schwartz, 13 (AMNH). Zion National Park, August 11, 1975, M. H. Sweet, 13 (TAMU). Zion National Park, August 13, 1929, P. W. Oman, R. H. Beamer, 63 (KU). Zion National Park, October 10, 1954, K. Goodarzy and G. F. Knowlton, 23 (KU). Wayne Co.: Caineville, June 2, 1966, G. F. Knowlton, 13 (UCD). Weber Co.: Ogden, June 10, 1965, G. F. Knowlton, 13 (USU). Ogden, May 27, 1954, G. F. Knowlton, 13 (KU). Plain City, July 19, 1940, G. F. Knowlton, 13 (USU). Virginia: Charlottes Co.: Charlottesville, July 5, 1928, L. C. Woodruff, 13 (KU). Washington: Benton Co.: Hanford Site ALE, T11N, R26E, sec. 35, 1200 ft, April 20, 1994, R. S. Zack, 23, 2♀ (WSU). Columbia Co. : 17 mi W of Pomeroy on Highway 12, July 7, 1979, G. Stonedahl, 123 (AMNH). Grant Co. : Coulee City, June 12, 1959, Madge and Kelton, Iva axillaris (Asteraceae) , 23 (CNC). 21 mi W of Moses Lake, May 30, 1957, E. F. Dailey ( Asteraceae ), 53, 5♀ (WSU). Wisconsin: Eau Claire Co.: Eau Claire, June 14, 1986, D. A. Rider, 63 (DAR). Waukesha Co.: Muskego, July 21, 1939, P. B. Lawson, 73 (KU). Wyoming: Goshen Co.: Lingle, May 17, 1955, W. D. Fronk (CNC). No specific locality, June 9, 1949, D. G. Denning (CNC). Torrington, September 13, 1955, W. D. Fronk (CNC). Laramie Co.: Albin, August 10, 1949, D. G. Denning (CNC). Natrona Co. : 9 mi E of Casper, August 12, 1962, J. G. and B. L. Rozen, 13 (AMNH). Niobrara Co. : 18 mi N of Lusk, August 10, 1971, J. G., B. L., and K. C. Rozen, 23 (AMNH). Park Co.: Powell, July 10, 1935, D. G. Denning (CNC). Platte Co.: Chugwater, August 30, 1954, W. D. Fronk (CNC). No specific locality, July 28, 1949, D. G. Denning (CNC). Wheatland, September 1, 1949, R. E. Pradt (CNC). Sheridan Co.: Bighorn Mountains, 6500 ft, August 17, 1927, H. H. Knight (CNC). Uinta Co. : Evanston, July 17, 1961, G. F. Knowlton, 13 (UCD). Lyman, June 25, 1935, J. Russell, 13 (KU).

Figures

Fig. 1. Habitus views of Chlamydatus spp.: C. artemisiae–C. opacus. See appendix 1 for localities.

Fig. 3. Coloration patterns on legs of Chlamydatus spp.

Fig. 4. Male genitalia of Chlamydatus spp.: C. artemisiae–C. keltoni. See appendix 1 for localities.

Fig. 7. Scanning electron micrographs of Chlamydatus associatus, 3, Ontario: St. Catharines. A. Head, frontal view. B. Head, lateral view. C. Setae on hemelytra. D. Mesothoracic spiracle and meta­