AMNHNOVITATES novi 00194 Mp_1 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3305, 69 pp., 23 figures, 1 table 13 December, 2000 Revision of the North American Plant Bug Genus Megalopsallus Knight, with the Description of Eight New Species from the West (Heteroptera: Miridae: Phylinae) RANDALL T. SCHUH1 ABSTRACT Megalopsallus Knight is revised, with 29 valid species recognized; 8 species are described as new and 13 previously described species are newly treated as junior synonyms. Merino- capsus Knight is treated as a junior synonym of Megalopsallus. The species Europiella albi- pubscensKnightandEuropiellamonticolaKnight,formerlyplacedinMegalopsallusaretreat- ed as incertae sedis and belonging to Europiella, respectively. Habitus and male genitalic illustrations are provided for all Megalopsallus species; scanning micrographs of the head, scent-gland evaporatory area, vestiture, and pretarsus are included for selected species. Akey to males is presented. The majority of Megalopsallus spp. are recorded from dry interior portions of the American West; two species are recorded from the Gulf Coast and East Coast of the United States. Host information is presented for most species, indicating an obligate association with halophytes, most in the families Ephedraceae (Ephedrales) and Chenopodi- aceae and Solanaceae (Angiospermae). The relationships of Megalopsallus within the Phylini are discussed. INTRODUCTION the type. Later, Knight (1968) described nu- merous species in the genus Europiella Reu- Knight (1927) described Megalopsallus, with five included species, designating atri- ter,manyofwhichweresubsequentlymoved plicis Knight from the Texas Gulf Coast as to Megalopsallus by Schuh et al. (1995), on 1GeorgeWillettCuratorofEntomologyandChair,DivisionofInvertebrateZoology,AmericanMuseumofNatural History. Copyright(cid:113)AmericanMuseumofNaturalHistory2000 ISSN0003-0082/Price$9.60 AMNHNOVITATES novi 00194 Mp_2 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 2 AMERICAN MUSEUM NOVITATES NO. 3305 the basis of male genitalic structure. The lat- mentsforallspeciesaregivenintable1,and ter authors did not, however,dealwithissues these data should be used for making size of synonymy and description of new taxa comparisons. All measurements are given in within Megalopsallus. millimeters. Species treatmentsarepresented Knight (1968) described Merinocapsus, in alphabetical order. with a single included species. Schuh (1986) later described an additional new species in Megalopsallus Knight Merinocapsus and treated Ankylotylus Megalopsallus Knight, 1927: 224 (n. gen.). Knight, 1968, with a single includedspecies, as a junior synonym. Merinocapsus Knight, 1968: 34 (n. gen.). NEW SYNONYMY. The present paper provides a detailed re- Ankylotylus Knight, 1968: 55 (n. gen.; syn. by vision of Megalopsallus, with a total of 29 Schuh, 1986: 217). valid species of these relatively colorful bugs; three of those species were previously TYPE SPECIES: Megalopsallus atriplicis placed in Merinocapsus Knight. Thirteen Knight, 1927. pre-existing names are treated as junior syn- DIAGNOSIS: Recognized by weakly flat- onyms, eight species are described as new, tend, lanceolate, silvery or white setae gen- habitus and genitalic illustrations are provid- erally distributed on dorsum and thoracic ed for all species, scanning micrographs are pleuron, intermixedwithpaleordark,reclin- provided for some structures from selected ing, simple setae (figs. 1C, 17D), elongate species, and a key to species is presented. nearly parallel-sided body form in males The relationships of Megalopsallus to other contrasted with ovate body form of females groups within the Phylini are discussed. (figs. 7–12), broad, usually short head in Although many of the species here placed males (figs. 7–12), and small, relativelysim- in Megalopsallus were originally described ple vesica of male genitalia (figs. 13–16) inEuropiella byKnight,primarilyontheba- with apex blunt or as a single or bifid, short sis of pretarsal structure (Knight, 1968), the apical projection, and gonopore sclerite morphology of the male genitalia and other (when present) lacking barbs. Coloration of- structures offers no suggestion of a close re- ten largely red or green, a few species cas- lationship between the two groups. The pul- taneous, but then appendages often partly to villi show considerable size variation in Me- entirely red. Known species feeding on hal- galopsallus, similar to that seen in Atracto- ophytes, primarily members of Ephedraceae, tomus Fieber (Stonedahl, 1990), and are Chenopodiaceae, and Solanaceae. therefore of little use in diagnosing the Megalopsallus was confounded by Knight group. Contrary to Knight’s (1927) original (1968) with Europiella Reuter, but is easily characterization, the pulvilli (pseudarolia of distinguished by the more flattened body Knight) are not ‘‘entirely absent,’’ but range form in most species, the simpler, more del- from small and Plagiognathus-like (fig. 17) icate structure and much smaller size of the to greatly enlarged and coveringmuchofthe male genitalia, and the preference for halo- ventral claw surface (fig. 18). phytic hosts rather than members of the As- Hosts are documented for nearly all of the teraceae. Some dark-colored species, suchas included species. The massive number of knowltoni and nigrofemoratus are similar in available host records allows for plant-insect appearance to some Plagiognathus species, associations to be critically assessed. It is but are always separable by the form of the now clear on which plant species most Me- male genitalia; furthermore, most Plagiog- galopsallus species breed, as well as on nathus spp. have only simple setae on the which plant species they do not breed. Al- dorsum and elsewhere on the body. The pul- though hostfidelity isnotabsoluteinallspe- villi are sometimes large and cover most of cies, it is shown to be very strong for most. the ventral claw surface in Megalopsallus The habitus photographs are not all repro- (figs. 18C, 20D, but see fig. 17E), whereas duced at a comparable scale. Thus, relative in Plagiognathus the pulvilli are always sizes of the taxa should not be assessed by small and cover only a small area near the comparing the figures. Detailed measure- base of the claw. As noted above, Knight’s AMNHNOVITATES novi 00194 Mp_3 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 2000 SCHUH: REVISION OF MEGALOPSALLUS 3 (1927) characterization of pulvillar structure often sclerotized and attenuated, as either a in Megalopsallus was in error. single spine or bifid, sometimes membra- REDESCRIPTION:Male:Smalltomoderately nous; (secondary) gonopore either apical or large species, total length 2.31–5.20, length subapical, occasionally only weakly sclero- apex clypeus–cuneal fracture 1.73–3.31, tized; gonopore scleritesubtendinggonopore width across pronotum 0.83–1.32; usually often present and well sclerotized (figs. 13– elongateandmoreorlessparallelsided(figs. 16, gs), never with barbs as in many Atrac- 7–12). COLORATION AND VESTITURE: totomus species; paramere and phallotheca Coloration, including all appendages, rang- typical of Phylini, without distinctive char- ing from totally pale, white, to almosttotally acteristics (figs. 14–16). black, often greenish or reddish. Vestiture of Female: Total length 2.28–3.98, length dorsum comprising reclining or recumbent, apex clypeus–cuneal fracture 1.74–2.96, pale to dark, simple setae intermixed with width across pronotum 0.96–1.42; often white, silvery, or rarely somewhat golden, broadly oval to elongate ovoid and of more weakly to moderately flattened, usually lan- robustbodyformthanmales(figs.7–12);an- ceolate,sometimeswoolly,setae(figs.7–12), tennal segment2 of slightlysmallerdiameter latter type also occurring on thoracicpleuron thaninmales,sometimesspindle-shaped(fig. and abdominal venter. Tibial spines pale or 23). dark, with or without dark spots at bases. DISCUSSION: Comparison of the 29 species STRUCTURE: Body form generally some- here recognized within Megalopsallus indi- what flattened; head broad, short longitudi- cates that the continued treatment of Meri- nally and clypeus not visible from above. nocapsus as a separate group almost certain- Eyes sometimes bulging and removed from ly renders the former group paraphyletic. pronotal margin, more commonly conform- Therefore, Merinocapsus is treated as a ju- ing to curvature of frons and to anterior nior synonym of Megalopsallus. pronotal margin. Antennal segment 2 cylin- The affinities of Megalopsallus appear to drical (figs. 5, 6) or more rarely spindle- be with Atractotomus Fieber, Knightomiro- shaped, especially in females (fig. 23). Hem- ides Stonedahl and Schwartz, Phoenicocoris elytraoftenelongatetostronglyelongateand Reuter, Pinomiris Stonedahl and Schwartz, nearly parallel-sided, abdomen often reach- and possibly Chlamydatus Curtis, based par- ing only to cuneal fracture, less frequently ticularly on the male genitalic structure, the hemelytra not so elongate and body form genitalia being relatively small, the vesica elongate ovoid. Metathoracic scent-gland being formed of a single strap, and the sub- evaporatory area as in figs. 1B, 3B, etc.;me- apical secondary gonopore frequently being sothoracic spiracle with an elongate, con- subtended by a gonopore sclerite (Kelton, spicuous area of ‘‘mushroom bodies’’dorsad 1965; Stonedahl, 1990; Stonedahl and of spiracle (figs. 3B, 4B). Claws ranging Schwartz, 1996). Although several Megal- from relatively short and curving only near opsallus species have the gonopore sclerite apex (fig. 1D) to relatively long and slender, (figs. 13–16, gs) described by Stonedahl smoothly curving over entire length (fig. (1990), none have barbs on the sclerite. The 17E); pulvilli ranging from minute (fig.17E) apex of the vesica in Megalopsallus is usu- to large and covering nearly entire ventral ally attenuated, in theformof asingle,short, claw surface (fig. 18C). Abdomen in males simple spine, or isbifid.Atractotomus,Chla- flattened dorsoventrally, broad basally and mydatus, and Phoenicocoris are Holarctic in broadly curving and narrowing toward rela- distribution, whereas the other genera are re- tively small genital capsule (fig. 17B). stricted to the Nearctic. MALE GENITALIA: Genital capsule and In addition to the above described mor- genitalia small relative to size of animal(fig. phological differences, the host associations 17B); vesica formed of a single strapusually in Megalopsallus are distinctive within this twisted to form an S shape (figs. 13, atripli- related group of genera. Megalopsallus spp. cis), more rarely forming a weak coil (fig. are restricted to halophytes in the genus 13, brendae) or not twisted and in the shape Ephedra of the primitive seed-plant group of a J (figs. 16, punctatus); apex of vesica Ephedrales and among the angiosperms pri- AMNHNOVITATES novi 00194 Mp_4 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 4 AMERICAN MUSEUM NOVITATES NO. 3305 marily to several genera in the Chenopodi- is moderately to greatly elongateandclothed aceae and to the solanaceous genus Lycium. with long to very long more-or-less erectse- Knightomiroides, Phoenicocoris, Pinomiris, tae. The vesica appears to be formed of a and some Atractotomus spp. feed on the single strap, as in Megalopsallus, and the Coniferales with the remainingAtractotomus genital capsule is relatively small. The gen- spp. breeding on a variety of angiosperms; eral appearance and coloration of Nasocoris none are known to feed on the Chenopodi- tesquorum Kerzhner are similar to those of aceae, Solanaceae, or Ephedraceae (Stone- Megalopsallus pallidus and the dorsal vesti- dahl, 1990; Stonedahl and Schwartz, 1996). ture is also similar to that of Megalopsallus A very few other Nearctic Phylini feedon spp. (Wagner, 1973). halophytes. Among those that do is Tanner- Psallomimus Wagner, with one species in ocoris sarcobati Knight, 1970, which is Egyptandtherestofitsspeciesfurthersouth found only on Sarcobatus, but which shows in Africa (Schuh, 1995), has relatively small no obvious relationship with Megalopsallus genitalia and the vesica is formed of a single onthebasisofhabitusormalegenitalicmor- strap as in Megalopsallus. However, theves- phology. titure is always of simple setae, and thebugs The Holarctic species Atomoscelisonustus are otherwise Plagiognathus-like in appear- (Fieber),1861,belongingtoanotherwisePa- ance. The only known host record is from learctic group, apparently feeds exclusively Solanum sp. (Linnavuori, 1993). on ruderal chenopods; most Nearctic records Psallopsis Reuter has15describedspecies of this species are as Atomoscelis modestus from the Mediterranean, Middle East, and (VanDuzee),1914[seeKerzhnerandSchuh, Central Asia; all breed on members of the 1998, for synonymy with Atomoscelis onus- Chenopodiaceae (Schuh, 1995). Although tus (Fieber)]. The vesica of Atomoscelis ap- host preferences in Psallopsis might suggest pears always to be formed of a single strap a relationship with Megalopsallus, the male basally, but is divided in two at the base of genitalia indicate otherwise, the vesicabeing the secondary gonopore, a condition never formed of two chitinous straps, rather than seen in the species here assigned to Megal- one as is the case in Megalopsallus(Wagner, opsallus. Furthermore, Atomoscelis spp. are 1975). usually ovate and much less strongly sexu- In conclusion, it appears that Megalopsal- ally dimorphic than Megalopsallus spp. lus as here diagnosed is restricted to the Ne- A few other Palearctic Phylini merit ex- arctic, and within that region to halophytic amination as possible Megalopsallus rela- plant groups. My conception of Megalopsal- tives on the basis of morphology and/or host lus is relatively broad, because when treated preferences.Theyarediscussedherewiththe otherwise the result would be several more objective of determining whether Megalop- difficult to diagnose genera. sallus as a monophyletic group extends be- AlthoughMegalopsallusshowsitsgreatest yond the Nearctic. species diversity in saline environments in Camptotylidea Wagner comprises 28 spe- interior western North America, it also oc- cies (Konstantinov, 1999) occurring primar- curs in the eastern United States—primarily ily in desert areas; many of them feed on along the coastline—from Texas to Connect- chenopods.ThevestitureofCamptotylideais icut, and most recently has been recorded composedofonlysimplesetaeandthevesica from coastal halophytes in the Dominican is always extended apically well beyond the Republic. Unfortunately, eastern North secondary gonopore, unlike the conditions American is relatively poorly collected and found in all species here assigned to Megal- few host records are available, in stark con- opsallus. trast to the West where nearly all species Nasocoris Reuter from the Mediterranean have well-documented hosts. and adjacent areas appearsto feedexclusive- The following key to species is designed ly on Ephedra (Schuh, 1995), which might for use with male specimens. For most spe- suggest a relationship with Megalopsallus. cies the description of coloration will also However, the clypeus is distinctively flat- pertain tofemales,althoughtable1willhave tened and extended and antennal segment 1 to be consulted for measurements. For a few AMNHNOVITATES novi 00194 Mp_5 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 2000 SCHUH: REVISION OF MEGALOPSALLUS 5 species it will be necessary to dissect the sarcobati Knight, 1969 (Europiella) male genitalia to be confident of the identi- schwartzi, new species fication. Sometimes host information will al- sparsus Van Duzee, 1918 (Europiella) low accurate identification of speciessuchas stitti Knight, 1968 (Europiella) humeralis and nigrofemoratus, which on the suaedae Knight, 1925 (Psallus) basis of external morphology and coloration teretis, new species may be virtually indistinguishable. viridiventris Knight, 1968 (Europiella) KEY TO MALES OF MEGALOPSALLUS KNIGHT CHECKLIST OF SPECIES NAMES PROPOSED OR USED WITHIN MEGALOPSALLUS KNIGHT 1. Dorsum, including head, nearly unicolorous pale, usually greenish, sometimes almost adustus Knight, 1927 (Megalopsallus) white or yellowish or pinkish; dorsum albipubescens Knight (Europiella) [incertae sometimes with small reddish spotsoroth- sedis] er markings on head, pronotum, and scu- arizonae Knight (Megalopsallus) tellum, or light brown spots at bases of se- atriplicis Knight, 1927 (Megalopsallus) tae ............................. 2 – At least head, and often much of dorsum, atriplicis Knight, 1968 (Psallus) [preoccu- darkened, ranging from pale reddish to pied; see punctatus] nearly black ..................... 15 balli Knight, 1968 (Europiella) 2. Membrane marmorate; entire dorsum, venter, brendae, new species and appendages pale green (fig. 7); legs brevicornis Knight, 1968 (Europiella) with some small brown spots; male geni- brittoni Knight, 1927 (Megalopsallus) talia as in figure 13 (San Joaquin Valley, californicus, new species California; host unknown) .. californicus diversipes Knight (Europiella) – Membrane not marmorate; coloration of dor- ephedrae Knight, 1968 (Merinocapsus) sum pale, sometimes greenish or pinkish ephedrellus, new species ................................ 3 3. Antennalsegment1almostentirelydark;dor- femoralis Kelton, 1980 (Megalopsallus) sum pale green (see fig. 7, brendae) ... 4 flammeus, new species – Antennal segment 1 entirely pale; dorsum franseriae Knight, 1969 (Europiella) greenish or otherwisepale,althoughsome- froeschneri Schuh, 1986 (Merinocapsus) times with reddish or brownish on prono- grayiae Knight, 1968 (Europiella) tum and scutellum (see fig. 10, pallidus) humeralis Van Duzee, 1923 (Sthenarus) ................................ 5 knowltoni Knight, 1970 (Europiella) 4. Femora pale green with some distinct black latifrons Knight (Europiella) spots (fig. 7); vertex and calli sometimes lycii Knight, 1968 (Europiella) weakly infuscate; frons often with a dark marmoratus Knight, 1968 (Megalopsallus) transverse line at level of base of clypeus; genitaliaasinfigure13(MojaveDesert;ex montanae Knight, 1968 (Europiella) Lycium, Chenopodiaceae sp.) .. brendae monticola Knight (Europiella) (see Euro- – Femora castaneous, usually appearing some- piella) what mottled (fig. 11); head and pronotum multipunctipes Knight, 1970 (Europiella) with some small red spots; genitalia as in nicholi Knight, 1968 (Europiella) figure 16 (Mojave Desert and south;exAl- nigricaput, new species lenrolfea, Salsola, Suaeda) ..... pictipes nigrofemoratus Knight, 1968 (Europiella) 5. Body and appendages entirely pale (see fig. nuperus Van Duzee, 1923 (Oncotylus) 10, pallidus) ...................... 6 pallidus Knight, 1968 (Nevadocoris) – At least some areas not pale, e.g., medial ar- pallipes Knight, 1968 (Ankylotylus) eas of pronotum and scutellum, thoracic sternum, spots on femora, and dark bases parapunctipes, new species of tibial spines (see fig. 12, sparsus) .. 7 pictipes Van Duzee, 1918 (Plagiognathus) 6. Eyes white (fig. 10); larger species, total punctatus, new name length 4.01–4.35, length apex clypeus–cu- punctipes Knight, 1968 (Europiella) neal fracture 2.61–2.83, width acrosspron- rubricornis Knight, 1968 (Europiella) otum 1.09–1.19; genitalia as in figure 15 rubropictipes Knight, 1927 (Megalopsallus) (Nevada; ex Atriplex, Grayia) ........ rufiventris Knight, 1968 (Europiella) ........................... pallidus AMNHNOVITATES novi 00194 Mp_6 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 6 AMERICAN MUSEUM NOVITATES NO. 3305 – Eyes red (fig. 12); smaller species, total atmostthoracicsternumdarkened,thoracic length 3.46–3.70, length apex clypeus–cu- pleuron and abdomen pale; setae on dor- neal fracture 2.26–2.37, width acrosspron- sum sometimes with brown bases .... 12 otum 0.96–1.03; genitalia as in figure 16 12. Dorsum pale, nearly white, more or less uni- (Great Basin; ex Sarcobatus) schwartzi formlycoveredwithtinybrownspotsatba- 7. Eyes red or reddish, distinctly protuberant, ses of setae on a lighter background (fig. especially in males (fig.12,rubropictipes); 11); large broad-bodied, total length 3.78– femora often with some reddish spots or 4.15, length apex clypeus–cuneal fracture markings ........................ 8 2.47–2.70, width across pronotum 1.16– – Eyes pale (fig. 12, sarcobati), grayish, or 1.27;genitaliaasinfigure16(extremewest- blackish(fig.12,sparsus),neverbrightred ern Texas, New Mexico, Arizona, southern or strongly protruding; femora spotted, but Nevada; ex Atriplex) ........ punctatus never with red ................... 11 – Dorsum, although uniformly pale, neverwith 8. Dorsum largely pale, cream colored, prono- brown spots atbases ofsetae;sizevariable tum (except humeralangles)andscutellum ............................... 13 contrastingly reddish or brownish (fig.10); 13. Elongate slender species, total length 3.28– genitalia as in figure 15 (southern Nevada 4.12, length apex clypeus–cuneal fracture and Utah; ex Ephedra) ........ pallipes 1.93–3.16, width of head 0.65–0.74, width – Dorsum unicolorous or nearly so, at most across pronotum 0.91–1.00; coloration with some small red spots on head and pale; ex Atriplex ................. 14 pronotum ........................ 9 – Shorter, more robust species, total length 9. Relativelysmall,slenderspecies(fig.7),total 3.04–3.26, length apex clypeus–cuneal length 3.25–3.77, length apex clypeus–cu- fracture 2.05–2.22, width of head 0.84– neal fracture 2.20–2.48, head relatively 0.87, width across pronotum 1.01–1.08; narrow, width across eyes 0.77–0.84; gen- coloration always at least weakly greenish italiaasinfigure13(InteriorwesternNorth (fig. 12); genitalia as in figure 16 (Great America, western Great Plains south to Basin; ex Sarcobatus) ....... sarcobati southern Texas; ex Atriplex, Sarcobatus) 14. Thoracic sternum infuscate; genitalia as in .......................... atriplicis figure 15; total length 3.28–3.72, length – Larger, more robust species, total length at apex clypeus–cuneal fracture 2.12–2.42, least3.10,lengthapexclypeus–cunealfrac- width of head 0.65–0.73, width across ture at least 2.27, width across eyes 0.85– pronotum 0.91–1.00 (fig. 10) (SW Great 0.94 ........................... 10 Basin; ex Atriplex confertifolia) ..... 10. Femora usually distinctly reddish, in contrast ...................... parapunctipes to greenish coloration of dorsum (fig. 12); – Thoracic sternum never dark; genitalia as in males elongate, females sometimesweakly figure 16; total length 2.91–4.12, length to strongly brachypterous; genitalia as in apex clypeus–cuneal fracture 1.93–3.16, figure 16 (Saskatchewan south to New width of head 0.73–0.74, width across Mexico and west to Oregon; ex Atriplex, pronotum 0.92–0.99 (fig. 11) (Great Basin Salicornia, Suaeda) ...... rubropictipes andsouthernRockyMountains;exAtripex) – Femora, if reddish, usually not noticeably ......................... punctipes contrasting with remaining coloration (fig. 15. Membrane marmorate; dorsum usually tan 10); sexual dimorphism weak to nearlyab- with a greater or lesser amount of reddish sent(fig.10);genitaliaasinfigure15(Tex- to brown spotting; all femora strongly red- as Gulf coast north to Colorado and Utah, dishbrown(fig.9);genitaliaasinfigure15 coastal Sinaloa, Mexico, Dominican Re- (Arizona and adjacent areas; ex Allenrol- public;exAtriplex,Batis,Salicornia,Suae- fea, Salicornia) .......... marmoratus da) ........................ nuperus – Membrane not marmorate; dorsum some- 11. Eyes black or blackish (fig. 12); thoracic times variably reddish brown with brown pleuron and venter and abdominal venter spots at bases of setae on lighter back- obviously infuscate; at least head often ground and femora reddish (fig.7,brittoni, with dark markings, sometimes also ante- femoralis) ....................... 16 rior pronotal lobe and scutellum (fig. 12); 16. General coloration, including legs and veins genitalia as in figure 16 (Saskatchewan ofmembrane,intenselyandcompletelyred south to Texas and west to southern Cali- or reddish (fig. 7, 8) .............. 17 fornia; ex Atriplex) ........... sparsus – General coloration variable, but never com- – Eyespale,usuallywhiteorpalegray(fig.12, pletely reddish ................... 18 sarcobati);headneverwithdarkmarkings; 17. Pronotum and scutellum sometimes darker AMNHNOVITATES novi 00194 Mp_7 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 2000 SCHUH: REVISION OF MEGALOPSALLUS 7 thanremainderofdorsum(fig.7);genitalia part or all of scutellum, and extreme base asinfigure14(centralNevada;exSarcob- of hemelyta ranging from pale red to cas- atus baileyi) ............... flammeus taneous, in contrast to white or pale green – Pronotumandscutellumofsameshadeasre- remainder of dorsum .............. 24 mainder of dorsum (fig. 8); genitalia as in – Coloration variable, but never exactly as figure 14 (southern Utah to northern Baja above, and if hemelytra mostly pale then California; ex Ephedra) ....... ephedrae abdomen mostly dark, or at most partially 18. Coloration of dorsum entirely dark, ranging palewithatleastsomereddishorinfuscate from brown to black (figs. 9, 10, 12, 21, areas ........................... 25 23) ............................ 19 24. Coloration as in figure 11; abdomen pale – Coloration of dorsum never completely dark, green, strongly contrasting withcastaneous rangingfromcompletelypaletopartlydark thoracic pleuron and venter; genitalia as in ............................... 22 figure 16 (Great Basin; ex Sarcobatus) 19. Relatively large species, total length 4.51– ........................ rubricornis 5.20, length apex clypeus–cuneal fracture – Coloration as in figure 8; abdomen pale 2.79–3.31, width across pronotum 1.15– green, thoracic pleuron and venter pale or- 1.32; antennal segment 2 slender, of uni- ange; genitalia as in figure 13 (westTexas; form diameter, never spindle-shaped (fig. ex Ephedra) .............. ephedrellus 9);genitaliaasinfigure14(northernGreat 25. Head, pronotum, and usually scutellum en- Basin; ex Sarcobatus vermiculatus) .... tirely castaneous to nearly black, or if not ..........................knowltoni entirelyso,thenonlyposteriorhumeralan- – Smaller species, total length in males 2.57– gles of pronotum and sometimes apex of 3.88, length apex clypeus–cuneal fracture scutellum pale ................... 26 1.74–2.46, width across pronotum 0.94– – Head, pronotum, and scutellum not entirely 1.16; shape of antennal segment 2 straight castaneous, often largely pale ....... 30 or spindle-shaped ................. 20 26. Darkcolorationofpronotumcontrastingwith 20. Antennal segment2inmalesenlarged,cylin- that of hemelytra, the latter ranging from drical,butdiameteratmostslightlygreater largely pale to distinctly orange or reddish than diameter of segment 1 (figs. 12, 22); ............................... 27 antennal segment 2 in females moderately – Dark coloration of pronotum usually not to strongly spindle-shaped (figs. 12, 23); strongly contrasting with coloration of antennaeandlegsconspicuouslyreddishin hemelytra; if hemelytra pale, then cuneus most specimens; genitalia as in figure 16 never reddish .................... 29 (southernNevadatoZacatecas,Mexico;ex 27. Head,pronotum,entireventer,andallfemora Lycium) ..................... teretis deep reddish to castaneous, contrasting – Antennalsegment2cylindricalinbothsexes, with pale (but not white) coloration of neverspindle-shaped,diameterslightlyless hemelytra(figs.5,9);cuneusalwaysweak- thanthatofsegment1;vesicanotasinfig- ly reddish; genitalia as in figure 15 (south- ure 16; coloration of legs and antennae ern Arizona, Mexico; ex Lycium) ..... sometimes reddish ................ 21 ............................nicholi 21. Genitalia as in figure 15; antennal segments – Head, pronotum, and scutellum nearly black; 1 and 2 varying from pale to reddish (fig. hemelytra, including cuneus, deep orange 10)(CanadatocentralMexico;exAtriplex, to red orange .................... 28 Grayia, ) ............. nigrofemoratus 28. Head, pronotum, and scutellum dull; femora – Genitalia as in figure 14; antennal segments unicolorous dark (fig. 8); tibiae infuscate; 1 and 2 varying from pale to dark (fig. 9) genitalia as in figure 14 (southern Nevada (southern Nevada and California, Arizona, and Utah south to central Baja California; Baja California; ex Lycium) .. humeralis ex Ephedra) ................ ephedrae 22. Head and eyes castaneous, strongly contrast- – Head, pronotum, and scutellum polished and ingwithpalegreentonearlywhiteremain- moderately to strongly shining (fig. 8); derofdorsum(figs.6,9);antennalsegment femora pale at least on distal one-half; tib- 1, thoracic venter (including coxae), and iae pale (southern Nevada, Utah, and Cal- much of femora also castaneous; genitalia ifornia; ex Ephedra) ....... froeschneri asinfigure15(Arizona,MojaveDesert;ex 29. Genitalia as in figure 15; antennal segments Lycium) .................. nigricaput 1 and 2 usually pale, although sometimes – If head and eyes castaneous, then remainder reddish (fig. 10) (Canada to central Mexi- of dorsum never entirely pale ....... 23 co; ex Atriplex, Grayia, and Sarcobatus) 23. Head, pronotum, mesoscutum, sometimes ..................... nigrofemoratus AMNHNOVITATES novi 00194 Mp_8 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 8 AMERICAN MUSEUM NOVITATES NO. 3305 – Genitalia as in figure 14; antennal segments cate; tibiae pale with dark spots at bases of 1 and 2 usually dark, less commonly red- pale spines. SURFACE AND VESTITURE: dish or pale (fig. 9) (southern Nevada and Dorsum smooth, weakly shining, clothed California,Arizona,BajaCalifornia;exLy- with recumbent, pale, simple setae inter- cium) .................... humeralis mixed with silvery, weakly flattened setae. 30. Hemelytramostfrequentlyentirelypale,usu- STRUCTURE: Hemelytra moderately elon- ally weakly greenish or grayish; head, an- gate, nearly parallel sided; eyes in males terior lobe of pronotum, mesoscutum, and more rarely hemelytra broadly dark (fig. large, protuberant (fig. 7); labium reaching 12); genitalia as in figure 16 (Texas to posteriormarginofmiddletrochanters;claws southern California; ex Atriplex) ..... relatively long and slender, smoothly curv- ........................... sparsus ing; pulvilli minute. MALE GENITALIA: – Coloration of dorsum reddish, often spotted Vesica weakly sigmoid, slender (fig. 13), at- or mottled, never greenish or grayish 31 tenuated apically; gonopore small,subapical, 31. Larger species, total length 3.57–3.82, length not subtended by a gonopore sclerite (fig. apex clypeus–cuneal fracture 2.58–2.71, 13). width across pronotum 1.18–1.29 (fig. 7); Female: Total length 2.97–3.19, length antennal segment 2 relatively long, length apex clypeus–cuneal fracture 2.14–2.33, 1.04–1.16;genitaliaasinfigure13(coastal Connecticut south to Florida and east to width across pronotum 0.99–1.08; relatively Texas); host unknown ......... brittoni stout, ovoid (fig. 7). – Smaller species, total length 3.09–3.24, HOSTS: Atriplex matamorensis, Atriplex length apex clypeus–cuneal fracture 2.16– texana, Atriplex sp., Sarcobatus vermicula- 2.29, width across pronotum 0.99–1.07 tus, S. baileyi (Chenopodiaceae). (fig.7);antennalsegment2relativelyshort, DISTRIBUTION: Interior of western North length 0.83–0.92; genitalia as in figure 13 America east to the western plains south to (northern Rockies; ex Salicornia) .... southern Texas. ......................... femoralis DISCUSSION: In the present paper atriplicis is conceived as widespread and not strongly Megalopsallus atriplicis Knight hostspecific.Thisapproachmightberevised Figures 7, 13 under more critical analysis. My treatmentis Megalopsallus atriplicis Knight, 1927: 224 (n. influenced by the fact that genitalia cannot sp.). be used to readily recognize this species, or DIAGNOSIS: Recognized by relativelysmall to distinguish it from nuperus and rubropic- size, protuberant red eyes, and distinctly red tipes, in particular. Also, nearly all material spotsonthehead,pronotum,andfemora(fig. from the eastern part of the range, including 7). Possibly most easily confused with the type locality, was from older collections, schwartzi and nuperus. Former species also which made comparisons of coloration dif- occurring on Sarcobatus at same localities, ficult. Finally, a vast section of the western but lacking red spots and protuberant eyes; plains and Rocky Mountains was virtually latter species more robust, without red spots unrepresented in the available sample of ma- and distinctive sexual dimorphism of atripli- terial. cis. Megalopsallus femoralis, marmoratus, SPECIMENS EXAMINED: USA. — Arizona: and pictipes frequently with red spots, but Coconino Co.: 1 mi E of Tuba City on Rt. without distinctly red eyes and strong sexual 163, 5000 ft, June 16, 1983, R. T. Schuh and dimorphism. M. D. Schwartz, Sarcobatus vermiculatus DESCRIPTION: Male: Small, elongate, total (Chenopodiaceae), 4(cid:63), 3(cid:47) (AMNH). Cali- length 3.25–3.77, length apex clypeus–cu- fornia: Inyo Co.: 2 mi E of Big Pine, June neal fracture 2.20–2.48, width across pron- 10, 1976–June 22, 1978, J. D.Pinto,Sarcob- otum 0.95–1.04. COLORATION: General atusvermiculatus(Chenopodiaceae),2(cid:63),5(cid:47) coloration cream to faded orange;vertex,an- (UCR). Mono Co.: 8 mi W of Nevada state terior margin of pronotum, and all femora line on Rt. 359, 6700 ft, July 2, 1983, R. T. with some reddish spots (fig. 7); eyes red; Schuh and M. D. Schwartz, Sarcobatus ver- thoracic pleuron and venter, all coxae, and miculatus (Chenopodiaceae), 6(cid:63), 12(cid:47) much of abdomen in females heavily infus- (AMNH). Colorado: Mineral Co.: Creede, AMNHNOVITATES novi 00194 Mp_9 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 2000 SCHUH: REVISION OF MEGALOPSALLUS 9 June21,1990,J.T.andD.A.Polhemus,1(cid:63), 16, 1979, M. J. Smith, N. Cobb, 1(cid:47) (OSU). 2(cid:47) (JTP). Moffat Co.: GatesofLodore,June T41S R35E S5, July 6, 1979, M. J. Smith, 28, 1979, D. A. Polhemus, 1(cid:63), 1(cid:47) (JTP). N. Cobb, 1(cid:63) (OSU). T36S R35E S8, June Rio Blanco Co.: W Evacuation Creek, 4 mi 26, 1979–September 4, 1979, M. J. Smith SE of state line on Rt. 45, 6400 ft, July 9, and N. Cobb, 5(cid:63), 1(cid:47) (OSU). T34S R35E 1981, M. D. Schwartz, Sarcobatus vermicu- S10 NE, July 12, 1979, Neil Cobb, 1(cid:47) latus (Chenopodiaceae), 1(cid:47) (AMNH).Mon- (OSU). Texas: Calhoun Co.: Port Lavaca, tana: Carbon Co.: Bear Creek between Red August 19, 1925–September 10, 1925, 7(cid:63), Lodge and Belfry, 5000 ft, August 12, 1986, 4(cid:47) (USNM). Cameron Co.: Brownsville, Schwartz and Stonedahl, Sarcobatus vermi- April 17, 1925, Div. Cotton Insects, Atriplex culatus (Chenopodiaceae), 2(cid:63), 13(cid:47) texana (Chenopodiaceae), 2(cid:63) (USNM). (AMNH). Nevada: Elko Co.: Elko, July 12, Brownsville, April 25, 1926, T. C. Barber, 1965, H. H.Knight,2(cid:63),17(cid:47) (USNM).Utah Atriplex matamorensis (Chenopodiaceae), state line on Utah Rt. 30, 4760 ft, June 25, Paratypes: 7(cid:63), 14(cid:47) (CNC, USNM); holo- 1983, Schuh and Schwartz, Sarcobatus ver- type: male (USNM). Brownsville, February miculatus (Chenopodiaceae), 4(cid:63), 27(cid:47) 1, 1926, T. C. Barber, Atriplex matamorensis (AMNH). Esmeralda Co.: 13 mi W of Lida (Chenopodiaceae), Paratypes: 7(cid:63), 10(cid:47) on Rt. 3, 1938 m, July 13, 1980, R. T. Schuh (USNM).Brownsville,February15,1925,T. and G. M. Stonedahl, Sarcobatus vermicu- C. Barber, Atriplex matamorensis (Cheno- latus (Chenopodiaceae), 1(cid:63), 1(cid:47) (AMNH). podiaceae), Paratypes: 1(cid:63) (USNM). Lander Co.: 11 mi S of Rt. 50 on Rt. 376, Brownsville, June 23, 1908, 1(cid:63), 3(cid:47) T17N R44E, 5800 ft, June 28, 1983, Schuh (USNM); Paratypes: 6(cid:63), (USNM). Browns- and Schwartz, Sarcobatus vermiculatus ville, May 1, 1925, Div. Cotton Insect, Atri- (Chenopodiaceae), 1(cid:63), 14(cid:47) (AMNH). 7.5 plex matamorensis (Chenopodiaceae), 15(cid:63), mi S of Rt 50 on Rt 376, T17N R44E, 5900 15(cid:47) (USNM). Utah: Box Elder Co.: S. Va- ft, June 28, 1983, R. T. Schuh and M. D. led Site, August 24, 1972, W. J. Hanson, 1(cid:63) Schwartz, Sarcobatus vermiculatus (Cheno- (USU). Cache Co.: Logan, July 22, 1938, G. podiaceae), 1(cid:47) (AMNH). Lincoln Co.: Ca- F. Knowlton, D. E. Hardy,1(cid:47)(USU).Emery thedral Gorge State Park, June 17, 1986, J. Co.: Green River, May 21, 1963, G. F. B. Knight and K. R. Helms, 8(cid:63), 1(cid:47) Knowlton, 1(cid:63) (USU). Garfield Co.: 14.3 S (AMNH). Nye Co.: 2.5 mi NE of Gabbs off of Rt. 95 on Rt. 276 (3.4 mi N of Starr Rt. 844, 4800 ft, July 2, 1983, R. T. Schuh Spring turnoff), 5000 ft, June 19, 1983, and M. D. Schwartz, 8(cid:63) (AMNH). 30 mi S Schuh and Schwartz, 10(cid:63), 15(cid:47) (AMNH). of Rt. 50 on Rt 376, T13N R14E, 6000 ft, Grand Co.: E of Moab, Colorado River June 30, 1983, R. T. Schuh and M. D. bridge, May 26, 1979, J. T. and D. A. Pol- Schwartz, Sarcobatus vermiculatus var. bail- hemus, 5(cid:63), 7(cid:47) (JTP). San Juan Co.: 25 mi eyi (Chenopodiaceae), 1(cid:63) (AMNH). 35 mi N of Monticello on Rt.191, 5700 ft,July18, N of Tonapah, Coyote Hole Spring/Sevier 1986, R. T. Schuh, Sarcobatus vermiculatus Reservoir, T8 R42E S11 & 23, 6000 ft, June (Chenopodiaceae), 2(cid:47) (AMNH). SevierCo.: 29, 1983, R. T. Schuh and M. D. Schwartz, Richfield, July 15, 1929, E. W. Davis, 7(cid:63), 83(cid:63) (AMNH). 35 mi N of Tonapah, Coyote 3(cid:47) (USNM). Salina, July 2, 1943, G. F. Hole Spring/Sevier Reservoir, T8 R42E S11 Knowlton, 2(cid:63) (KU). Uintah Co.: 5–10 mi & 23, 6000 ft, June 30, 1983, Schuh, SE of Bonanza, 5000–5600 ft, May 14, Schwartz, Sarcobatus vermiculatus var. bail- 1982–September 6, 1982, M. D. Schwartz, eyi (Chenopodiaceae), 1(cid:63), 10(cid:47) (AMNH). Sarcobatus vermiculatus (Chenopodiaceae), Washoe Co.: 2 mi E of Vya on Rt. 8A, 1750 11(cid:63),16(cid:47)(AMNH).Bonanza,atWhiteRiver m, July 2, 1979, R. T.Schuh and B.M.Mas- Shale Project Trailers, T9S R24E Sec 23, sie, 2(cid:63), 26(cid:47) (AMNH). New Mexico: Eddy 5800 ft, May 6, 1982–July 7, 1981, M. D. Co.: April 24, 1979, Burke, Dolorme, Schwartz, Sarcobatus vermiculatus (Cheno- Schaffner, 5(cid:63) (TAMU). Taos Co.: Ojo Cal- podiaceae), 7(cid:63), 64(cid:47) (AMNH). Weber Co.: iente,June6,1982,D.A.andJ.T.Polhemus, Slaterville, July 12, 1967, G. F. Knowlton Atriplex sp. (Chenopodiaceae), 2(cid:63) (JTP). and L. E. Frank, 1(cid:63) (OSU). Washington: Oregon: Harney Co.: T41S R36E S8, July Yakima Co.: Yakima, August 15, 1931, A. AMNHNOVITATES novi 00194 Mp_10 WednesdayNov21200101:19PM2000 Allen Press • DTPro System File # 01TQ 10 AMERICAN MUSEUM NOVITATES NO. 3305 Fig. 1. Megalopsallus brendae, male, scanning micrographs. A. Lateral view of head and thorax. B. Mesothoracic spiracle and metathoracic scent gland evaporatory area. C. Detail of setae comprising dorsal vestiture. D. Lateral view of pretarsus. R. Rolfs, 11(cid:63), 10(cid:47) (USNM). Yakima, June somewhat heavier and more nearly forming 30, 1932, A. R. Rolfs, 10(cid:63), 8(cid:47) (USNM). a coil than in most other Megalopsallus spe- Wyoming: Sheridan Co.: Arvada, July 31, cies (fig. 13). 1927,H.H.Knight,Sarcobatusvermiculatus DESCRIPTION: Male: Relatively small, total (Chenopodiaceae), 1(cid:63), 2(cid:47) (USNM). length 2.74–3.16, length apex clypeus–cu- neal fracture 2.74–3.16, width across pron- Megalopsallus brendae, new species otum 0.90–1.01. COLORATION: Generally Figures 1, 7, 13 pale with a greenish tinge (fig. 7); eyes, transverse line on frons at level of base of HOLOTYPE: Male, California: San Bernar- clypeus, maxillary plate, antennal segment 1 dino Co.: 1.3 mi. S of Goffs, 845 meters, May 16, 1978, R. T. Schuh, ex ? Chenopo- (except base and apex), thoracic pleuron and dium sp. (Chenopodiaceae). Deposited inthe venter, and abdomen including genital cap- American Museum of Natural History. sule brownish black; femora with darkspots, DIAGNOSIS: Recognized by generally pale tibial spines black with dark bases. SUR- greenish coloration of dorsum contrasting FACE AND VESTITURE: Dorsum weakly with dark antennal segment 1; possibly most polished and moderately shining, covered easily confused with sarcobati on basis of with dark, recumbent, simple setae inter- pale green coloration; differing from that mixed with silvery, weakly flattened setae species in having antennal segment 1 dark (fig.1C);tibialspinesblackwithblackbases. andventerdarkenedinmales.Vesicainmale STRUCTURE:Moderatelyelongateandpar-