Noviltatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

NoviltatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3166, 15 pp., 27 figures May 23, 1996

Two New Genera for Pine-Inhabiting Species of Phylini in North America (Heteroptera: Miridae: Phylinae)

GARY M. STONEDAHL1 AND MICHAEL D. SCHWARTZ2

ABSTRACT Two new genera of phyline Miridae are de- P. monophyllae. The male genitalic structures of scribed from western North America to accom- all species are illustrated, and scanning electron modate taxa removed from existing genera and micrographs are presented for the head, dorsal several newly recognized species. Knightomiroides vestiture, femoral spines, pretarsus, and genital comprises the type species Psallus nigrovirgatus capsule ofrepresentative species. Habitus illustra- Knight and two new species, K. contortae and K. tions are provided for K. nigrovirgatus and P. ponderosae, and Pinomiris comprises the type spe- knighti. cies Lepidopsalluspini Knight and the new species

INTRODUCTION Among some pine-inhabiting Phylini from of this material now allows for the descrip- western North America examined for the re- tion of two new genera to accommodate one vision of the genus Atractotomus Fieber species removed from both Phoenicocoris (Stonedahl, 1990) were specimens that did Reuter and Psallus Fieber, plus three newly not fit within the revised diagnoses of that recognized species. genus or Phoenicocoris Reuter. Our analysis

1 Research Associate, Department ofEntomology, American Museum ofNatural History; Senior Scientific Officer, International Institute of Entomology, London. 2 Biological Resources Division, Centre for Land and Biological Resources Research, Ottawa.

sity ofCalifornia, Berkeley (UCB); T. J. Hen- ry, National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). We also thank G. duHeaume, In- ternational Institute of Entomology, London (IIE) for the dorsal habitus illustrations of Knightomiroides nigrovirgatus and Pinomiris knighti, and I. Boler (IIE) for assisting with the preparation of the scanning electron micrographs. Thomas J. Henry and Randall T. Schuh provided useful comments on an ear- lier version of the manuscript.

SYSTEMATICS

I f, IN Pinomiris, new genus .'N- (Figures 1-12, 26) TYPE SPECIES: Lepidopsallus pini Knight. DiAGNoSIS: Recognized by the dorsum with dense covering of moderately broad, silvery /11A ,X') 'S~~~~~~~~/1 white, scalelike setae, intermixed with heavy, dark, simple setae (fig. 3); pointed head with weakly produced, strongly sloping frons and tylus (fig. 2); length ofhead greater than length ofpronotum in dorsal view; labium reaching well beyond apices ofmetacoxae; dorsodistal surface of metafemora with 6-8 irregularly placed heavy spinules, these about half as Fig. 1. Dorsal habitus of Pinomiris knighti, d. long as surrounding bristlelike setae (figs. 5, 6); and by the structure of the male genitalia, especially the distally swollen left margin of the genital capsule (fig. 7) and robust vesica ACKNOWLEDGMENTS with nonspinose membranous lobe distal to In addition to material from the Centre for secondary gonopore (figs. 7, 9, 12). Land and Biological Resources Research, Ag- DESCRIPTION: Male. Macropterous; length riculture Canada, Ottawa (CNC), we thank from apex of tylus to cuneal fracture 1.58- the following individuals and institutions for 1.79 (all measurements in millimeters); width the loan ofspecimens: R. T. Schuh, American across humeral angles ofpronotum 0.74-0.91; Museum of Natural History, New York general coloration reddish brown to fuscous; (AMNH); J. T. Polhemus, Polhemus Collec- dorsal vestiture composed of moderately tion, Englewood, Colorado (JTP); A. Slater, broad, silvery white, scalelike setae inter- Snow Entomological Museum, University of mixed with stout, dark, simple setae (figs. 1, Kansas, Lawrence (KU); J. D. Lattin, Sys- 3). Head: Pointed anteriorly (fig. 2); frons and tematic Entomology Laboratory, Depart- tylus weakly produced, strongly sloping; pos- ment of Entomology, Oregon State Univer- terior margin weakly concave, conforming to sity, Corvallis (OSU); J. C. Schaffner, De- anterior margin of pronotum; vertex nearly partment of Entomology, Texas A&M Uni- flat medially, slightly less than half as broad versity, College Station (TA&M); F. G. as width of head across eyes; eye occupying Werner and C. Olsen, Department of Ento- most of head height in lateral view; genae mology, University of Arizona, Tucson about as broad as diameter of antennal seg- (UAZ); J. A. Chemsak, Department of En- ment I; buccal cavity ovate; gula narrow; la- tomology, California Insect Survey, Univer- bium reaching beyond apices of metacoxae. 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 3

Antennae: Inserted at level ofventral margin Female: Macropterous; length from apex of eye; segment I not quite reaching to apex of tylus to cuneal fracture 1.61-1.75; similar of tylus; segment II linear, about two-thirds to male in general appearance, except slightly as long as width of head across eyes; com- more ovoid, with shorter hemelytral mem- bined lengths of segments III and IV about brane, and with narrower, weakly clavate sec- equal to length of segment II; all segments ond antennal segment. with golden brown to dark brown, recum- DISTRIBUTION: Figure 26. Southern Great bent, simple setae; segment I with several Basin including parts of Arizona, Colorado, erect, stout bristles distally. Thorax: Prono- Nevada, New Mexico, and Utah. tum trapezoidal, more than twice as broad ETYMOLOGY: Named for the strict associ- as long, weakly convex, anterior and poste- ation of the included species with pinyon rior angles broadly rounded, lateral and pos- pines. terior margins nearly straight, anterior mar- DISCUSSION: Pinomiris knighti was origi- gin weakly convex; calli indistinct; mesoscu- nally described in the genus Lepidopsallus by tum broadly exposed; scutellum weakly ele- Knight (1968), a placement based primarily vated, nearly flat dorsally; thoracic pleurites on the characteristic dorsal vestiture of the and lateral regions ofabdominal sternites with bug. Stonedahl (1990) synonymized Lepi- dense covering of scalelike setae. Hemelytra: dopsallus with Atractotomus, transferring the Weakly rounded laterally, reaching well be- pine-inhabiting species of the former genus yond apex of genital capsule; embolium in- to Phoenicocoris. Our more detailed exami- distinct distally; cuneal incisure and fracture nation of the Holarctic genus Phoenicocoris well developed; cuneus about as long as broad. has revealed that knighti is not congeneric Legs: Femora about three times as long as with the type species, modestus (Meyer-Diir). broad; dorsodistal surface ofmetafemora with We are here erecting the new genus Pinomiris 6-8 irregularly placed strong spinules, these to accommodate knighti and its sister species about half as long as surrounding setae (figs. monophyllae. 5, 6); tibiae cylindrical, with three rows of heavy black spines; area ofinsertion ofspines on tibiae not darkened; tarsal segments I and Pinomiris knighti II subequal in length, segment III noticeably (Kerzhner and Schuh), longer; pretarsal claws moderately curved new combination distally; pulvilli well developed, covering (Figures 1-7, 11, 12, 26) about three-fourths of ventral claw surface; Lepidopsallus pini Knight, 1968: 51, 53 (n. sp., setiform parempodia relatively short and key); Henry and Wheeler, 1988: 470 (cat.). stout (fig. 4). Genitalia: Genital capsule: Dis- Phoenicocorispini: Stonedahl, 1990: 61 (n. comb.). tal surface adjacent to base of left paramere Phoenicocoris knighti: Kerzhner and Schuh, 1995: with broad, rounded swelling (fig. 7). Right 3 (n. name for Lepidopsallus pini Knight-ju- paramere: Simple, ovate, abruptly narrowed nior secondary homonym ofAtractotomus pini distally, apex acute. Left paramere: Anterior Fieber); Schuh 1995: 374 (cat.). process strongly developed; outer dorsal sur- DiAGNOsIs: Recognized by the characters face set with heavy setae (figs. 8, 11). Phal- given in the generic diagnosis; reliably dis- lotheca: S-shaped; broadly opened posteri- tinguished from P. monophyllae only by the orly; apical curved region broadly developed structure of the male genitalia, especially the (fig. 10). Vesica: Composed of single, broad, vesica with shorter apical process and non- sclerotized strap, strongly twisted medially; spinose gonopore sclerite (fig. 12) and left distal, upright region of strap more or less paramere with anterior process shorter than perpendicular to base; gonopore subapical, posterior process (fig. 1 1). with thick walls visible in lateral view, sub- DESCRIPTION: Male. Length from apex of tended by well-developed sclerite, the latter tylus to cuneal fracture 1.55-1.79; width suspended in membrane and sometimes set across humeral angles ofpronotum 0.74-0.86; with surface spines; vesica distal to gonopore reddish brown or dark brown general color- with distinctly protruding membranous lobe ation. Head: Brown to dark reddish brown, (figs. 9, 12). mandibular plate sometimes paler yellowish 4 AMERICAN MUSEUM NOVITATES NO. 3166

II_ Ii

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Figs. 2-7. Scanning electron micrographs of Pinomiris knighti. 2. Lateral view of head. 3. Scalelike setae on hemelytra. 4. Pretarsus. 5. Dorsal view of metafemur, with detail of dorsodistal surface. 6. Dorsodistal surface of metafemur. 7. Posterolateral view of genital capsule showing vesica protruding from phallotheca. 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 5 brown; width across eyes 0.61-0.66; width of paramere with anterior and posterior pro- vertex 0.27-0.31; antennal coloration as de- cesses more or less evenly developed (fig. 8). scribed for P. monophyllae, segment II some- DESCRIPTION: Male. Length from apex of times narrowly darkened at base and apex; tylus to cuneal fracture 1.58-1.77; width length of antennal segment II 0.36-0.5 1; la- across humeral angles ofpronotum 0.84-0.91; bium reaching to base of genital capsule, reddish brown or dark brown general color- length 1.09-1.24. Thorax: Uniformly brown ation, hemelytra sometimes brownish gray. or dark brown, lateral angles of mesoscutum Head: Yellowish brown to dark brown; tylus, and apex of scutellum sometimes yellowish especially apically, usually darker than sur- brown. Hemelytra: Uniformly brown or dark rounding sclerites; width across eyes 0.63- brown; base of corium sometimes yellowish 0.68; width ofvertex 0.27-0.30; antennal seg- brown; hemelytral membrane evenly suf- ment I dark brown to fuscous; segment II pale fused with fuscous, veins slightly darker than yellowish brown, length 0.47-0.53; segments surrounding membrane. Legs: Color as de- III and IV brown or yellowish brown; labium scribed for P. monophyllae; tibiae sometimes reaching beyond apices of metacoxae to near narrowly darkened basally. Genitalia: Fig- base of genital capsule, length 1.13-1.20. ures 11, 12. Thorax: Uniformly brown to fuscous, me- DISTRIBUTION: Figure 26. Northern Ari- soscutum sometimes slightly paler, especially zona, western and southern Colorado and on lateral angles. Hemelytra: Brown or dark New Mexico; inhabits Pinus edulis. brown, usually lightly tinged with red; corium MATERLAL EXAMINED: Arizona: Coconino sometimes brownish gray, especially basally Co.: 40 mi S Grand Canyon, 3.VILL.1917, H. and on distal third; cuneus grayish brown to H. Knight (USNM, Knight Coll.) 12; Wil- fuscous; hemelytral membrane lightly and liams, 7000 ft, 24.VI.1925, A. A. Nichol evenly suffused with fuscous, veins slightly (CNC, USNM, Knight Coll.) 26 (paratypes) darker than membrane. Legs: Femora red- Colorado: Costilla Co.: Ft. Garland, dish brown to fuscous, narrowly pale apically; lO.VIII.1925, ex Pinus edulis, H. H. Knight- tibiae and tarsi pale yellowish brown, apex 16 (USNM, holotype), 16 (CNC, paratype), of tarsal segment III and pretarsus dark 52 (USNM, allotype, paratypes). Montrose brown. Genitalia: Figures 8-10. Co.: 10 mi W Montrose, 8.VlI.1980, ex Pinus DISTRIBUTION: Figure 26. Currently known edulis, J. T. & D. A. Polhemus (JTP) 56, 1 12. only from the basin range region of Nevada. Rio Blanco Co.: W Evacuation Creek, 4 mi ETYMOLOGY: Named for its host plant, Pi- SE Colorado St. Border on UT St. Rt. 45, nus monophylla. 6400 ft, 9.VII.1981, ex Pinus edulis, M. D. TYPES: Holotype 6. Nevada, WhitePine Co., Schwartz (AMNH) 12. New Mexico: Otero 4.2 mi W of Baker, 2031 m, 14.VII.1980, ex Co.: Cloudcroft, 23-25.VII.1967, L. A. Kel- Pinus monophylla, R. T. Schuh & G. M. ton (CNC) 16; 2 mi SW Mayhill, 17.VII.1979, Stonedahl (AMNH). Paratypes. Nevada: Delorme, McHugh & Schaffner (TA&M) 86, Clark Co.: Charleston Peak, 20.VII. 1982, ex 172; 6 mi N Pinon, 12.VIII.1982, ex Pinus Pinus edulis [misidentification for Pinus edulis, J. C. Schaffner (TA&M) 16, 102. So- monophylla], J. T. Polhemus (JTP) 36, 72. corro Co.: 12 mi W Magdalena, 15.VIII.1982, White Pine Co.: same data as holotype ex Pinus edulis, J. C. Schaffner (TA&M) 12. (AMNH) 46, 72. ADDITIONAL MATERIAL EXAMINED: Neva- da: Nye Co.: Nevada test Site, Area 12M(W), 25.VII.1962, ex Pinus monophylla (USNM, Pinomiris monophyllae, new species Knight Coll., paratype ofLepidopsallus pini) (Figures 26) 12. Lander Co.: 7 mi SE of Austin, 8-10, 20.VII.1961, J. F. Lawrence (UCB) 16, 12. DIAGNOSIS: Distinguished from P. knighti Churchill Co.: reststop [on Rt 2] W ofCarroll by the structure of the male genitalia, es- Summit, T16N R37E, 6500 ft, 5.VIII.1982, pecially the more robust vesica with spines ex Pinus monophylla, M. D. Schwartz on the gonopore sclerite (fig. 9) and the left (AMNH) 22. 6 AMERICAN MUSEUM NOVITATES NO. 3166

Figs. 8-12. Male genitalia of Pinomiris species. 8-10. Pinomiris monophyllae. 8. Left paramere. 9. Vesica. 10. Phallotheca. 11, 12. Pinomiris knighti. 11. Left paramere. 12. Vesica.

Knightomiroides, new genus narrow, apically acute, scalelike setae, inter- (Figures 13-25, 27) mixed with long, dark, simple setae (figs. 13, 15); pretarsus with large pulvilli, reaching TYPE SPECIES: Psallus nigrovirgatus Knight. nearly to apex of claw (fig. 16); metafemora DIAGNOSIS: Recognized by the head with without spinules on dorsodistal surface (fig. broad, weakly inflated mandibular plate and 17); and by the structure ofthe male genitalia, well-developed genae (fig. 14); usually with especially the large genital capsule, robust shiny black tylus contrasting with pale head phallotheca (fig. 21), anterior process of left (ifhead mostly darkened then tylus and man- paramere with a long subapical seta, and ve- dibular plate more shiny than remainder of sica with small field of spines on membra- head); dorsal vestiture composed ofrelatively nous lobe distal to gonopore, and elongate, 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 7 weakly spinous gonopore sclerite (figs. 20, 23, femora about four times as long as broad; 25). dorsodistal surface of metafemora without DESCRIPTION: Male. Macropterous; length spinules (fig. 17); tibiae cylindrical, with three from apex of tylus to cuneal fracture 1.81- rows of heavy black spines; area of insertion 2.19; width across humeral angles of pro- of spines on tibiae narrowly infuscated; pre- notum 0.84-1.03; pale yellowish brown to tarsal claws straight dorsally, moderately dark brown general coloration; dorsal vesti- curved distally; pulvilli large, covering most ture composed of narrow, adpressed, silvery of ventral surface of claw; setiform parem- white, scalelike setae intermixed with long, podia relatively short and stout (fig. 16). Gen- dark, suberect, simple setae (figs. 13, 15). italia: Genital Capsule: Large, broadly pro- Head (fig. 14): Weakly rounded anteriorly; duced distally. Rightparamere: Elongo-ovate, moderately produced anterior to antennal with distinct apical process. Left paramere: fossae in lateral view; posterior margin nearly Anterior process relatively short, with long straight, conforming to anterior margin of subapical seta; posterior process narrow; out- pronotum; vertex very weakly convex, about er dorsal surface set with heavy setae (figs. half as broad as width of head across eyes; 19, 22, 24). Phallotheca: S-shaped; broadly frons weakly rounded, junction with tylus in- opened posteriorly; distal region beyond bend distinct; tylus weakly produced; mandibular broadly developed (fig. 21). Vesica: Com- plate broadly produced, noticeably inflated; posed oflong, thin, sclerotized strap, strongly eyes occupying about three-fourths of head twisted medially; distal upright portion of height in lateral view; genae well developed, strap very long, more or less perpendicular about twice as broad as diameter ofantennal to base, apex very narrowly attenuate; gon- segment I; buccal cavity large, subspherical; opore subapical with aperture visible in lat- gula nearly obsolete; labium reaching to api- eral view, walls moderately thick, subtended ces of mesocoxae or slightly beyond. Anten- by elongate sclerite reaching to lowest point nae: Inserted at level ofventral margin ofeye; of basal angle; gonopore sclerite suspended segment I reaching to or slightly beyond apex in membrane, set with fine spines at least on of tylus; segment II linear, length less than basal third adjacent to gonopore; vesica distal width of head across eyes; combined lengths to gonopore with weakly to moderately pro- of segments III and IV about equal to length truding membranous lobe, usually set with of segment II; all segments with golden to series of very fine spines (figs. 20, 23, 25). brown, reclining simple setae, segments II and Female: Macropterous; length from apex IV usually with scattered paler setae; segment of tylus to cuneal fracture 1.61-1.75; similar I with several erect, heavy, black bristles. to male in general appearance, except slightly Thorax: Pronotum trapezoidal, slightly more ovoid, with shorter hemelytral mem- greater than twice as broad as long, weakly brane, and with narrower, weakly clavate sec- convex, very shallowly depressed between and ond antennal segment. behind calli; anterior and posterior angles of DISTRIBUTION: Figure 27. Western United pronotum broadly rounded, lateral margin States and southern British Columbia, Can- nearly straight, posterior margin broadly and ada. shallowly concave medially; calli weakly dif- ETYMOLOGY: Named in honor of the late ferentiated; mesoscutum broadly exposed; Harry H. Knight, whose scientific contribu- scutellum weakly elevated, nearly flat dor- tions greatly advanced our understanding of sally; thoracic pleurites and lateral areas of plant bug taxonomy, particularly in North abdominal sternites covered with scalelike America. setae. Hemelytra: Very weakly convex lat- DISCUSSION: The new genus Knightomi- erally, reaching well beyond apex of genital roides is here erected to accommodate Psallus capsule; embolium of even thickness nigrovirgatus and two new species of plant throughout; cuneal incisure and fracture dis- bugs distributed in western North America. tinct; cuneus slightly longer than broad; Psallus nigrovirgatus is clearly not congeneric membrane lightly suffused with fuscous, veins with the European type ofthe genus, P. roseus mostly pale. Legs: Pale yellow to dark brown; (Fabricius), the two species differing widely 8 AMERICAN MUSEUM NOVITATES NO. 3166

of tylus to cuneal fracture 1.89-2.20; width across humeral angles ofpronotum 0.84-1.03; ix pale gray ground color with brown to fuscous markings, usually yielding distinct striped pattern (fig. 13). Head: Pale yellow; tylus shiny fuscous; vertex and maxillary plate usually infuscated but mandibular plate remaining pale; width across eyes 0.72-0.77; width of vertex 0.36-0.39; antennal segment I yellowish brown to fuscous, apex always narrowly pale; segment II yellowish brown, sometimes slightly darker distally, length 0.59-0.68; segments III and IV yellowish brown to dark brown; length of labium 0.93-1.02. Thorax: Pronotum pale yellow or grayish yellow anteriorly, posterior third to half fuscous; pro- pleuron mostly pale, posterior margin sometimes narrowly to broadly infuscated; mesoscutum yellow, sometimes tinged with red; scutellum grayish yellow to fuscous, paler specimens with at least lateral margins tinged with fuscous. Hemelytra: Pale gray ground color with margins of clavus, inner half of corium, embolium, and most ofcuneus brown to fuscous; some specimens more extensively darkened with only medial region of clavus, corium bordering radial vein, and paracu- Fig. 13. Dorsal habitus of Knightomiroides ni- neus pale; membrane lightly to moderately grovirgatus, 6. suffused with fuscous. Legs: Yellowish brown; last tarsal segment and sometimes apex of tibiae brown to fuscous; anterior surface of in external morphology and characters ofthe femora usually with dusky spots medially in male genitalia. roughly linear series. Genitalia: Figures 18, 21-23. Knightomiroides nigrovirgatus DISTRIBUTION: Figure 27. Mountainous (Knight), new combination regions of Colorado, Arizona, and north- (Figures 13-18, 21-23, 27) eastern Utah. SPECIMENS EXAMINED: Arizona: Apache Psallus nigrovirgatus Knight, 1930: 130, 131 (n. Co.: Eagar, Apache Nat. For., 12- sp.); Carvalho, 1958: 125 (cat.); Henry and 13.VIII.1967, L. A. Kelton (CNC) 16. Co- Wheeler, 1988: 494 (cat.); Schuh, 1995: 410 conino Co.: Kaibab Lk., Kaibab Nat. For., (cat.). 16.VIII.1967, L. A. Kelton (CNC) 12; San DiAGNosIs: Distinguished from K. contor- Francisco Mts., 9.VIII.1930, E. D. Ball tae and K. ponderosae by the distinctly bi- (USNM, Knight Coll.) 12; 18.VIII.1927, P. colored pronotum and hemelytra (fig. 13); pale A. Readio (KU) 12. Gila Co.: Workman Crk, yellow head with shiny black tylus-vertex Sierra Ancha Mts, 5000 ft, 9.VII. 1964, at UV and maxillary plate also usually darkened; light, J. Burger (UAZ) 36. Pima Co.: Mt. and by the structure of the male genitalia, Lemmon, Santa Catalina Mts, 3-4.VIII. 1967, particularly the lobal sclerite of the vesica ex Pinus ponderosa, L. A. Kelton (CNC) 116, with limited spines mostly restricted to basal 232. Colorado: Boulder Co.: Nederland, third adjacent to gonopore and large spinose 31.VIII. 1972, L. A. Kelton, ex Pinus sp. lobe dorsal to secondary gonopore (fig. 23). (CNC) 22. Jefferson Co.: Falcon Pk., Indian REDESCRIPrION: Male. Length from apex Hills, 7200 ft, 21.VIII.1986, ex Pinus pon- 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 9

Figs. 14-18. Scanning electron micrographs of Knightomiroides nigrovirgatus. 14. Lateral view of head. 15. Scalelike setae on hemelytra. 16. Pretarsus. 17. Dorsodistal surface of metafemur. 18. Pos- terolateral view of genital capsule showing left paramere and apex of phallotheca. derosa, R. T. Schuh & J. T. Polhemus T. Schuh & J. T. Polhemus (AMNH) 16; (AMNH) 16, 2Q; Pine, 23.VII. 1907, C. A. Hill Trinidad, Stonewall, 8500 ft, 7.VIII. 1925, ex (CNC) 16 (paratype). Las Animas Co.: Stone- Pinus ponderosa, H. H. Knight (USNM, wall, 8000 ft, 25.VII.1968, ex Pinus ponder- Knight Coll.) 1 Y (paratype). Larimer Co.: osa, L. A. Kelton (CNC) 116, 6Y; 3 mi N Chambers Lk., Roosevelt Nat. For., 9200 ft, Stonewall, 8100 ft, 18.VIII.1986, at light, R. 11 .VIII. 1968, ex Pinus ponderosa, L. A. Kel- 10 AMERICAN MUSEUM NOVITATES NO. 3166

Figs. 19-25. Male genitalia of Knightomiroides species. 19, 20. Knightomiroides contortae. 19. Left paramere. 20. Vesica. 21-23. Knightomiroides nigrovirgatus. 21. Phallotheca. 22. Left paramere. 23. Vesica. 24, 25. Knightomiroides ponderosae. 24. Left paramere. 25. Vesica. ton (CNC) 16, 49; Estes Park, 24.VIII.1925, Verde Nat. Park, 7000 ft, 17-18.VII. 1968, ex ex Pinus ponderosa, H. H. Knight (CNC; Pinus ponderosa, L. A. Kelton (CNC) 146, USNM, Knight Coll.) 36 (holotype and para- 109. Park Co.: Wilkerson Pass, Pike Nat. For., types); Mt. Park, Poudre Can., 5.IX. 1972, L. 28.VIII. 1968, ex Pinus ponderosa, L. A. Kel- A. Kelton (CNC) 19; Mt. Park, Roosevelt Nat. ton (CNC) 36, 59. New Mexico: Los Alamos For., 6600 ft, 6.VIII. 1968, L. A. Kelton (CNC) Co.: 15 mi W Los Alamos., 23.VIII. 1979, L. 36, 49; Pingree Park, 22.VIII. 1925, ex Pinus A. Kelton (CNC) 16. Utah: Daggett Co.: Uin- ponderosa, H. H. Knight (USNM, Knight tah Mts., 0.2 mi N jct. Rts 44 & 260, 9 mi S Coll.) 16 (paratype). La Plata Co.: Durango, Dutch John, T2N R22E, 7500 ft, 2.VIII. 1981, 19.VII. 1968, ex Pinus ponderosa, L. A. Kel- ex Pinus ponderosa, M. D. Schwartz (AMNH) ton (CNC) 36, 19. Montezuma Co.: Mesa 16. 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI I1I

Knightomiroides contortae, DIScuSSION: The general coloration of K. new species contortae is quite variable, ranging from yel- (Figures 19, 20, 27) lowish brown to dark brown, with teneral specimens often being very similar in exter- DIAGNOSIS: Distinguished from K. nigro- nal appearance to the usually much paler K. virgatus and K. ponderosae by the yellowish ponderosae. In California and Oregon where brown to dark brown general coloration; head these two species are broadly sympatric, it is mostly darkened, not pale, with strongly con- advisable to consult the structures ofthe male trasting, shiny, fuscous tylus; and by the genitalia for positive identification, particu- structure ofthe male genitalia, especially the larly if the host plant is not known. shorter anterior process of the left paramere TYPES: Holotype 6. Oregon, Union Co., 0.1 (fig. 19) and numerous, coarse spines on the mi NE Tollgate Shopping Center on Hwy. gonopore sclerite of the vesica (fig. 20). 204, 17.VIII.1979, ex Pinus contorta, M. D. DESCRIPTION: Male. Length from apex of Schwartz (AMNH). Paratypes. Oregon: Des- tylus to cuneal fracture 1.82-2.19; width chutes Co.: Three Creeks Meadow, 18 mi S across humeral angles ofpronotum 0.89-0.98; Sisters, 6300 ft, ex Pinus contorta, J. D. Lattin brown or dark brown general coloration, (OSU): 1.IX.1977,26, 19 and 19.IX.1979, 36, rarely yellowish brown. Head: Brown or dark 49; Three Creeks Meadow, ex Pinus contorta brown, sometimes with yellowish brown (AMNH): 27.IX.1978, M. D. Schwartz, 19 markings, especially on frons and vertex; ty- and 19.IX.1979, G. Stonedahl, 19. HoodRiv- lus and mandibular plate usually darker and er Co.: T3S R9E Sec. 18, 4300 ft, 13.IX.1979, more shiny than remainder of head; width ex Pinus contorta, J. D. Lattin (OSU) 16. Linn across eyes 0.69-0.74; width of vertex 0.34- Co.: Big Lake, 4650 ft, L.IX.1979, ex Pinus 0.39; antennal segment I yellowish brown to contorta, J. D. Lattin (OSU) 29. Union Co.: fuscous; segment II brown or yellowish brown, same data as holotype (AMNH) 116, 139. sometimes darker distally, length 0.54-0.66; ADDITIONAL SPECIMENS: California: Las- segments III and IV brown to fuscous; length sen Co.: Bogard Cpgd., 24 mi W jct. Hwys oflabium 0.84-1.02. Thorax: Pronotum uni- 36 & 44, 10.VIII. 1980, ex Pinus contorta, G. formly brown or dark brown, rarely yellowish Stonedahl (AMNH) 56, 89. Tuoloumne Co.: brown anterior to calli; mesoscutum yellow- Yosemite Nat. Park, Yosemite Crk Ranger ish brown, sometimes broadly suffused with Stn., 21.VII.1946, ex Pinus murryana [= Pi- fuscous medially, lateral angles always pale; nus contorta], R. L. Usinger (UCB) 29. Mono scutellum brown to fuscous. Hemelytra: Pale Co.: Mammoth Lk., 29.VII. 1940, D. E. Har- grayish brown to dark brown, corium distally dy (KU) 16, 29. Idaho: Benewah Co.: Picnic and cuneus sometimes yellowish brown; Area, 2 mi W Santa at jct. Rts. 3 & 6, 2800 membrane moderately suffused with fuscous. ft, 6.VIII. 1986, ex Picea engelmannii, Schuh, Legs: Brown or yellowish brown; femora Schwartz & Stonedahl (AMNH) 19. Oregon: sometimes with faint dark marks mostly along Klamath Co.: 1 mi W Crescent, 4500 ft, midline ofanterior surface; tibiae sometimes 17.VII.1979, ex Pinus contorta, M. D. narrowly dark brown apically; tarsal segment Schwartz (AMNH) 29. Lane Co.: Scott Lake, II usually fuscous. Genitalia: Figures 19, 20. 25 mi E Rainbow, 4859 ft, 29.VIII.1960, ex DISTRIBUTION: Figure 27. Widely distrib- Pinus contorta, J. D. Lattin (OSU) 49. Ne- uted in mountainous regions ofwestern North vada: Washoe Co.: Little Valley Research America from Mammoth Lakes in the Sierra Area, 4.5 mi SW Washoe, T16N R19E Sec. Nevada Mountains, north through the Ore- 2, 6200 ft, 4.VIII. 1982, ex Pinus contorta, M. gon Cascades to northern Idaho, and south D. Schwartz (AMNH) 46, 19. Utah: Duchesne in the Rocky Mountains to Yellowstone Na- Co.: 0.5 mi S Avintaquin Cmpgrd. on Rt. 33, tional Park and the Bighorn Range in Wyo- TIlS R1OE, 9000 ft, 9.VIII.1981, ex Pinus ming, and the Wasatch Range in northeastern flexilis, M. D. Schwartz (AMNH) 26,29. Wy- Utah. oming: Big Horn Co.: 27 mi W Burgess Jct. ETYMOLOGY: Named for its primary host on Rt. 14, 7700 ft, 12.VIII.1986, ex Pinus plant, Pinus contorta. flexilis, Schuh, Schwartz & Stonedahl 12 AMERICAN MUSEUM NOVITATES NO. 3166

(AMNH) 106, 279. Park Co.: Yellowstone brook, 23.VII.1959, ex Pinus ponderosa, L. Nat. Park, 5 mi S Tower Falls, 7350 ft, A. Kelton (CNC) 186, 302. USA. California: 17.IX.1980, ex Pinus contorta, J. D. Lattin Modoc Co.: 18 mi S Rt. 139 toward Lookout, (OSU) 12. 1420 m, 6.VII. 1979, R. T. & J. Schuh (AMNH) 16. Siskiyou Co.: 0.5 mi S Lava Beds Nat. Mon. toward Medicine Lake, 5300 Knightomiroides ponderosae, ft, 27.VII.1986, ex Pinus ponderosa, R. T. new species Schuh (AMNH) 56, 32. Oregon: same data as (Figures 24, 25, 27) holotype (AMNH) 116, 1 O. ADDITIONAL SPECIMENS: CANADA. Brit- DiAGNosIs: Recognized by the uniformly ish Columbia: Summerland, 2-11 .VII. 1974, pale dorsal coloration; head with tylus, max- L. A. Kelton (CNC) 16, 12. USA. California: illary plate, and mandibular plate shiny fus- Lassen Co.: Bogard Cpgd., 24 mi Wjct. Hwys cous, and vertex usually broadly infuscated; 36 & 44,1 0.VIII. 1980, ex Pinus ponderosa, and by the structure of the male genitalia, G. Stonedahl (AMNH) 22. Tuolumne Co.: especially the nearly straight apical region of Yosemite Nat. Park., Nevada Falls, the vesica (figs. 25). 29.VII.1946, R. L. Usinger (UCB) 12. DESCRIPTION: Male. Length from apex of tylus to cuneal fracture 1.84-2.15; width across humeral angles ofpronotum 0.96-1.00; pale yellowish brown general coloration. DISCUSSION Head: Pale yellow or grayish white; tylus, The relationship of Pinomiris and Knigh- maxillary plate, and mandibular plate shiny tomiroides to other genera of Phylini is not fuscous; vertex broadly infuscated; width well understood. Stonedahl (1990) included across eyes 0.77-0.82; width of vertex 0.37- a character discussion for a group of phyline 0.39; antennae pale yellow to yellowish genera defined by the presence of scalelike brown, segments III and IV sometimes darker setae and structure of the vesica. Based on brown; length of antennal segment II 0.58- that discussion, the new genera have certain 0.62; length of labium 0.82-0.98. Thorax: affinities to Atractotomus and Phoenicocoris. Pronotum grayish white, calli usually lightly The head structure of Pinomiris is very tinged with brown; posterior lobe ofpronotal similar to that of Phoenicocoris; both have a disk usually darker brownish yellow, some- strongly sloping frons with narrow genae. times lightly suffused with fuscous; mesoscu- However, the buccal cavity and the posterior tum brown or yellowish brown, sometimes margin of the head in the two genera differ. lightly tinged with red; scutellum yellowish The cavity ofPinomiris is ovoid and the pos- brown, sometimes suffused with fuscous. He- terior margin conforms to the anterior mar- melytra: Pale yellowish brown, embolium and gin of the pronotum. In Phoenicocoris the most of cuneus somewhat paler; membrane buccal cavity is weakly semispherical and the very lightly suffused with fuscous. Legs: Pale posterior margin ofthe head is mostly straight. yellow or brownish yellow; tarsal segment III The rounded and subvertical frons (in lateral brown; femora sometimes with faint dusky view), concave posterior margin of the head, marks along midline ofanterior surface. Gen- and spherical buccal cavity of Knightomi- italia: Figures 24, 25. roides are similar to those of Atractotomus DISTIUBUTION: Figure 27. Cascade Range species. and Sierra Nevada Mountains from Cran- Both new genera have "Type 1" scalelike brook, British Columbia, Canada, south to setae previously defined by Stonedahl (1990) Yosemite National Park in California. as narrow, moderately flattened, apically ETYMOLOGY: Named for its host plant, Pi- pointed, and with distally converging ridges. nus ponderosa. The broad, flattened, apically serrate setae TYPES: Holotype 6. USA, Oregon, Grant with pustulate ridges ("Type 2" setae) found Co., Malheur Nat. For., Funny Bug Basin, on Phoenicocoris and some Atractotomus are T17S R32E Sec. 19, 23.VII.1979, ex Pinus not present on Knightomiroides and Pinom- ponderosa, M. D. Schwartz (AMNH). Para- iris. types. CANADA. British Columbia: Cran- Pinomiris has an irregular row of6-8 stout 1 996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 13

* *

* Pinomiris pini * Pinomiris monophyllae

Fig. 26. Distribution of Pinomiris species. spinules on the dorsodistal surface ofthe me- The pretarsal structures of Pinomiris and tafemora as found on Phoenicocoris and most Knightomiroides are similar. Both genera have Atractotomus. These spinules are not found relatively short, stout setiform parempodia on Knightomiroides species. and large, well-developed pulvilli that cover 14 AMERICAN MUSEUM NOVITATES NO. 3166

* Knightomiroides contortae * Knightomiroides nigrovirgatus A Knightomiroides ponderosae

Fig. 27. Distribution of Knightomiroides species.

most or all of the ventral claw surface. This Phoenicocoris has long, apically spatulate is similar to the pretarsal structure ofthe min- parempodia and small pulvilli (Stonedahl, iatus group of Atractotomus. In contrast, 1990). 1996 STONEDAHL AND SCHWARTZ: NEW GENERA OF PHYLINI 15

The vesica of Pinomiris and Knightomi- of Knightomiroides. The process in Knigh- roides is a single, strongly twisted strap sub- tomiroides also has a long, subapical seta. Both tended by a long gonopore sclerite usually new genera have broad genital capsules and covered with spinules. The secondary gono- robust phallothecae in comparison to Atrac- pore is subapical with a small, distal mem- totomus and Phoenicocoris. branous lobe. This lobe is spinose in Knigh- The large pulvillus, spines on the gonopore tomiroides. In lateral view the orientation of sclerite (although this character is homopla- the gonopore (apparently due to the degree sious), and coiled vesica support a group con- of twisting of the vesical strap) differs be- taining Atractotomus, Knightomiroides, and tween the two genera: in Pinomiris the side Pinomiris. Several features indicate a sister- ofgonopore is visible; in Knightomiroides the group relationship between Pinomiris and opening of gonopore is visible. The single, Knightomiroides, including the short, stout twisted vesical strap, usually with a spinose parempodia, broad genital capsule, robust gonopore sclerite is similar to Atractotomus. phallotheca, and the membranous lobe bor- The vesical strap of Phoenicocoris is more dering the distal margin ofthe secondary gon- tightly coiled, apically bifurcate, and has a opore. These and other characters should be short gonopore sclerite without spinules. assessed over a much broader range of taxa The anterior processes of the left para- before the problem of generic relationships meres ofboth new genera are distinctive. The can be fully resolved. process in Pinomiris is long compared to that

REFERENCES Carvalho, J. C. M. 1968. Taxonomic review: Miridae of the Ne- 1958. Catalogue of the Miridae of the world. vada Test Site and the western United Part II. Subfamily Phylinae. Arq. Mus. States. Brigham Young Univ. Sci. Bull., Nac. Rio de Janeiro, 45: 1-216. Biol. Ser. 9(3): 1-282. Henry, T. J., and A. G. Wheeler, Jr. Schuh, R. T. 1988. Family Miridae. pp. 251-507. In T. J. 1995. Plant bugs of the world: (Insecta: Het- Henry and R. C. Froeschner (eds.), Cat- eroptera: Miridae): systematic catalog, alog ofthe Heteroptera, or true bugs, of distributions, host list, and bibliogra- Canada and the continental United phy. New York: N. Y. Entomological States. Leiden and New York: E. J. Brill. Society. 1329 pp. Kerzhner, I. M., and R. T. Schuh Stonedahl, G. M. 1995. Homonymy, synonymy, and new com- 1990. Revision and cladistic analysis of the binations in the Miridae (Heteroptera). Holarctic genus Atractotomus Fieber Am. Mus. Novitates 3137: 11 pp. (Heteroptera: Miridae: Phylinae). Bull. Knight, H. H. Am. Mus. Nat. Hist. 198: 1-88. 1930. New Species of Psallus Fieb. (Hemip- tera, Miridae). Can. Entomol. 62: 125- 131. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates, Bulletin, and Anthropological Papers published during the last five years are available free of charge. Address orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, N.Y. 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: [email protected]

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