Pseudopsallus Van Duzee, 1916b: 224

SCHWARTZ, MICHAEL D., 2005, Redescription of Strophopoda aprica Van Duzee and the Description of Two New Genera and Five New Species from the Southwestern United States and Northern Mexico (Heteroptera: Miridae), American Museum Novitates 3489 (1), pp. 1-24 : 14-22

Pseudopsallus Van Duzee, 1916b: 224 (n.gen.).

REVISED DIAGNOSIS: Stonedahl and Schwartz (1986, 1988) distinguished the genus from other Orthotylini based on the dorsal vestiture and male genitalic structure. The inclusion of greggii and tiquiliae in Pseudopsallus requires the following modifications to the generic diagnosis: tergal processes sometimes absent from anterodorsal margin of genital aperture ( daleae and tiquiliae ); basal sclerotized portion of the ductus seminis rarely without a pair of spiculae surrounding the left and right sides on the dorsal surface ( tiquiliae ); and right ventral spicula of the vesica rarely minute ( greggii ).

DISCUSSION: A medial interior flange and associated troughlike surface ventral to the flange of the right paramere is shared by species of Pseudopsallus and Presidomiris Stonedahl and Schwartz, 1988 (see Stonedahl and Schwartz 1988: 17). However the gray general coloration with dark fuscous to black markings on the head and pronotum with dark suffusion on the hemelytra (fig. 1) will distinguish greggii and tiquiliae from species of Presidiomiris , which are consistently unmarked and have orange yellow to yellow green overall coloration. The small body size and strongly produced anteocular portion of the head of the two new species are unique for Pseudopsallus , but the prepoderance of characters, especially the body coloration and overall features of the male genitalia, are persuasive enough to place them in the genus.

The new species are generally similar to P. angularis ( Uhler, 1894) and P. daleae (Knight, 1968) in the pale gray to pale brown overall coloration with distinct black to fuscous markings on the head, pronotum, venter, and legs (fig. 1). Although P. tulare Stonedahl and Schwartz, 1986 has the ground coloration pale greenish yellow, it has dark markings similar to the other four species. Portions of the male genitalia, specifically the prominent distal hump of the medial interior flange on the right paramere (fig. 7F– H) and deflexed left tergal process adjoining the genital aperture (fig. 7A, B) of greggii are similar to those of angularis ( Stonedahl and Schwartz 1986: figs. 67, 68, 70, 71) and tulare . The absence of tergal processes (fig. 9A) and the small dorsal lobe of the left paramere (fig. 9E) in tiquiliae are also features seen in daleae ( Stonedahl and Schwartz 1988: figs. 44, 47, 48).

The scalelike setae of greggii (fig. 6C) and tiquiliae (fig. 8C) are similar in structure to those of the other species of Pseudopsallus ( Stonedahl and Schwartz 1986: figs. 11–25, 1988: fig. 34A–C). All members of the genus have scalelike setae that are strongly flattened, moderately to broadly lanceolate, apically truncate or acuminate, and with converging or parallel ridges. Such setae were defined as type 2 by Stonedahl and Schwartz (1986).

The pretarsus of the new species have strongly curved claws that are distinctly broadened basally, small pulvilli, and typically orthotyline lamellate and apically convergent parempodia (figs. 6D, 8D). The structure of the mesothoracic spiracle and metepisternal scent efferent system in the new species (figs. 6E, 8E) are unremarkable except for the relatively reduced evaporative area of the mesothoracic spiracle when compared with other North American Orthotylini (e.g., Schwartz and Scudder 2003: figs. 4E, 6D, 9C, 11B, 13B).

The incorporation of greggii and tiquiliae into the cladistically derived phylogeny presented in Stonedahl and Schwartz (1986, 1988) is beyond the scope of this present paper, which is intended only to provide documentation for newly discovered taxa.

The inclusion of two more taxa raises the number of species in Pseudopsallus to 25. In the key to Pseudopsallus species by Stonedahl and Schwartz (1986), greggii and tiquiliae will key to couplet 3 with angularis as well as daleae and tulare (see Stonedahl and Schwartz 1988: 15). The following addition to that key will distinguish these five species:

1. Clypeus entirely black, sometimes female with bilaterally paired and medial fuscous stripes; anteocular portion of head strongly produced (figs. 6A, B, 8A, B); body size less than 3.80..................... 2

— Clypeus entirely pale or with pale ground col­ or and bilaterally paired and medial fuscous to black stripes; anteocular portion of head short or moderately produced (similar to Stonedahl and Schwartz 1986: figs. 5, 6, 8); body size greater than 4.00.......... 3

2. Antennal segment 1 black; male without tergal processes on dorsal aperture of genital segment......... tiquiliae , new species

— Antennal segment 1 pale with a black basal annulus; male with tergal process on left dorsal margin of genital segment aperture................ greggii , new species

3. Dark marking on head limited to small bilateral pair on vertex near eyes; dorsal vestiture with pale simple setae; male without tergal processes on dorsal aperture of genital segment.......... daleae (Knight)

— Dark marking on head extensive, with bilateral stripes or blotches on frons, vertex, maxillary plate, and clypeus; dorsal vestiture with black simple setae......... 4

4. Body length 4.66–5.03?, 5.02–5.60/; pale greenish yellow general coloration with limited infuscation on pronotal disk and hemelytra; anteocular portion of head produced; male genitalia distinctive (see Stonedahl and Schwartz 1988: 15, 16, figs. 50–55).. tulare Stonedahl and Schwartz

— Body length 5.30–6.30?, 5.50–6.20/; brownish yellow or brownish gray general coloration with extensive infuscation on hemelytra; anteocular portion of head not produced; male genitalia distinctive (see Stonedahl and Schwartz 1986: 13, figs. 65– 71)................ angularis (Uhler)

The new species utilize host plants of the genus Tiquilia Persoon [; Coldenia Linnaeus ] or coldenia and crinklemat and are thus the only members of Pseudopsallus to be associated with the Boraginaceae . The known hosts of the other species of this endemic North American plant bug genus are from the Asteraceae (eight spp.), Fabaceae (one sp.), Hydrophyllaceae (two spp.), Lamiaceae (one sp.), Nyctaginaceae (two spp.), and Onagraceae (six spp.) ( Stonedahl and Schwartz 1986, 1988).

Pseudopsallus greggii , new species

Figures 1 , 6 , 7

HOLOTYPE: Male: ‘‘ USA: Texas: Brewster

Co. [,] Big Bend National Pk.[,] Rosilla Mts.,

298349330; 1038159400[,] August 5, 1991 [,] J. C. Schaffner[;] Taken on Tiquilia greggii (T. & G.) Richardson [ Boraginaceae ,] det A. M. Powell 991[;] Natl. Park. Serv.[,] Big Bend N. P.[,] Accession #650 [blue label].’’ Deposited in the Texas A&M University Insect Collection.

DIAGNOSIS: Along with tiquiliae , recognized among the species of Pseudopsallus by the relatively short body length and strongly produced anteocular portion of the head; greggii is also distinguished by its pale antennal segment 1, conspicuous black subapical patch on dorsal surface of the femora, black labium, wider vertex (0.35–0.37?, 0.39 –0.44/; tiquiliae 0.30–0.33?, 0.33– 0.40/), more produced anteocular portion of the head (0.30–0.34?, 0.34–0.41/; tiquiliae 0.26–0.30?, 0.29–0.36/), and longer labium (1.85–2.30?, 1.93–2.48/; tiquiliae 1.50– 1.85?, 1.66–1.95/).

DESCRIPTION: Male: Small, subparallel­sid­ ed. MEASUREMENTS: Total length 3.34 (2.95–3.80), length apex clypeus­cuneal fracture 2.35 (2.18–2.60), maximum width across hemelytra 1.29 (1.14 –1.43); head width 0.71 (0.68–0.74), vertex width 0.35 (0.35–0.37), anteocular length 0.32 (0.30– 0.34), eye width 0.17 (0.15–0.18), eye height 0.36 (0.34–0.38); antennal length segment 1: 0.21 (0.20–0.23), 2: 1.09 (0.98–1.24), 3: 0.81 (0.65–0.88), 4: 0.36 (0.31–0.39); labium length 2.13 (1.85–2.30), reaching insertion of left paramere; pronotum width 1.06 (0.96– 1.16), pronotum length 0.57 (0.53–0.63). COLORATION (fig. 1): creamy gray; head with clypeus dark fuscous to black, dark markings on anterior portion of maxillary plate, extreme anterior margin of mandibular plate, two variable pairs of small bilateral marks on frons and one pair of spots on temporal areas, collum infuscate; antennal segment 2–4 dark fuscous to black; labial segment 1 black on ventral margin, segments 2– 4 black; black mark on dorsal margin of metepisternum; calli with diffuse fuscous to black markings, pronotal disk sometimes with diffuse orange brown spots; mesoscutum black, scutellum brown basomedially; portions of endocorium, paracuneus, and cuneus variably infuscate; membrane strongly infuscate, veins creamy white; abdomen with variable dark fuscous markings on ventral margin and laterally on subdorsal margin; tumid setose region on left side of genital aperture; parameres deeply fuscous; black spots on legs, femora with obvious black area subapically; tibiae black basally and distally; tarsi black. SURFACE AND VESTITURE: Subshining, smooth, dorsal vestiture of mixed, widely scattered, slightly expanded, silvery, scalelike setae (fig. 6C) and moderately densely distributed, pale, reclining simple setae; tibiae with pale reclining setae, dark brown spines, and parallel rows of minute black spicules. STRUCTURE: Head triangular in dorsal view, length from apex of clypeus to vertex 91% of head width across eyes; frons gently rounded; clypeus obvious in dorsal view; anteocular distance in lateral view equal to 3.53 diameter of antennal segment 1, anteocular distance 13% greater than distance posterior to antennal fossae in dorsal view; eye height in lateral view equal to 93% of head height; pronotum subquadrate in dorsal view; corial margins subparallel. GENI­ TALIA: Genital segment: aperture with one pair of subequal length, flattened left tergal processes, LM with three to four prominent apical serrations, LL with serrate lateral margin (figs. 6F, 7A, B); left surface of segment, between tergal processes and paramere insertion, tumid with stouter setae than on remainder of genital segment.(figs. 6F, 7A). Left paramere: DLL equally broad basally and medially, with four or five apical spines, basalmost spines somewhat recurved (figs. 6F, 7J); LAL gradually attenuate, apex rounded, slightly recurved, lateral surface with spinules (figs. 6F, 7I). Right paramere: MIF marginally serrate, with prominent, ventrally serrate, tubercle distally; troughlike region ventral to MIF with spinules (fig. 7F, G); DLR broadly truncate, smooth (fig. 7E); lateral surface slightly concave ventrally (figs. 6F, 7H). Phallotheca: aperture sinuate, right side with notch and concavity (fig. 7D). Vesica: DS prominent, smooth, slightly attenuate; RVS minute, situated ventrally; LVS long, smooth, strongly attenuate (fig. 7C).

Female: Macropterous. Similar to male in color, vestiture, and structure, except for smaller eyes, wider vertex, slightly shorter hemelytra, and sometimes pale clypeus with dark bilateral paired and medial stripe. MEA­ SUREMENTS: Total length 3.23 (2.95– 3.45), length apex clypeus­cuneal fracture 2.45 (2.20–2.63), maximum width across hemelytra 1.30 (1.17–1.43); head width 0.72 (0.70–0.75), vertex width 0.40 (0.39–0.44), anteocular length 0.38 (0.34 –0.41), eye width 0.15 (0.14–0.16), eye height 0.35 (0.33–0.36); antennal length segment 1: 0.22 (0.20–0.23), 2: 1.11 (0.97–1.20), 3: 0.83 (0.68–0.91), 4: 0.37 (0.31–0.41); labium length 2.31 (1.93–2.48), reaching ninth abdominal sternite; pronotum width 1.04 (0.94–1.11), pronotum length 0.54 (0.49– 0.60).

ETYMOLOGY: Named for the occurrence of all specimens on the host species Tiquilia greggii .

HOST: Tiquilia greggii (Torrey and A. Gray) A. T. Richardson , plume coldenia.

DISTRIBUTION: Southern New Mexico to southwestern Texas.

PARATYPES: USA: New Mexico: Eddy Co.: Campsite, 32821.49N 103846.99 W, D. R. Delorme, H. L. Carroll, ex light, 1? ( TAMU) ; Hope , July 22, 1967, L. A. Kelton, 1? ( CNC) . Hidalgo Co.: Granite Gap , 1 mi N of, August 30, 1974, J. D. Pinto 1?, 1/ ( UCR) . Otero Co.: Mountain Park , June 27, 1940, D. E. Hardy, 1? ( KU) ; White Sands , June 27, 1940, R. H. Beamer, D. E. Hardy, 3?, 1/ ( KU) . Texas: Brewster Co.: same label data as holotype, 26?, 22/ [fig. 1 adult dorsal habitus photographs] ( TAMU) ; Big Bend Ranch S. N. A., 1.8 mi N of McGuire’s Tanks, August 7, 1991, J. C. Schaffner, 3?, 1/ ( TAMU) ; Cottonwood Campground, Big Bend National Park , September 27–28, 1986, J. T. Doyen, ex black and white light, 2? ( UCB) ; Rosillos Mts. Lodge, Big Bend Natl. Pk., September 21, 1990, Zolnerowich and Cecora, ex black light, Z90/074, 4? ( TAMU) . Hudepeth Co.: 0.5 mi E of Indio Mountain Reserve Station , 308469240N 1058009480W, 4100 ft, April 12, 2002, Diaz and Gillogly, ex MV light, 2? ( TAMU) . Presidio Co.: Presidio : October 5, 1929, S. E. Jones, 5? ( TAMU) ; August 12, 1968, J. E. Hafernik, ex black light, 1? ( TAMU) ; 10 mi N of, August 18, 1965, J. C. Schaffner, 29?, 10/ ( TAMU) . 13 mi N of, (AMNH, CNC, USNM, TAMU): 14 July 1968, M. L. Allender, 2?, 6/. C. L. Cole , ex Tiquilia greggii det. A. M. Powell 1992: July 14, 1966, 4?, 10/; July 26, 1966, 3?, 7/; August 8, 1966, 1/; September 13, 1966, 14?, 24/; September 30, 1966, 3/. Shafter, 2 mi N of, J. E. Hafernik: July 2, 1968, 5?, 2/ ( TAMU) ; July 30, 1968, ex Tiquilia greggii det. A. M. Powell 1992, 7?, 5/ ( TAMU) ; August 9, 1968, 4/ ( TAMU) . Val Verde Co.: Seminole Canyon St. Pk, April 15, 1989, J. Heraty, 1? ( TAMU) . Webb Co.: Laredo , 27 mi S of, June 28, 1979, J. D. Pinto 1/ ( UCR) ; Mines Rd at San Ambrosia Creek , May 2, 1994, W. Godwin, ex black light, 2? ( TAMU) .

Pseudopsallus tiquiliae , new species

Figures 1 , 8 , 9

HOLOTYPE: Male: ‘‘ USA: California: Imperial Co. , 5.4 mi NW Ocotillo on rt S2, April 23, 1980, Schwartz & Russell, Tiquilia palmeri (A. Gray) A. Rich [ards.] ( Boraginaceae ).’’ Deposited in the American Museum of Natural History.

DIAGNOSIS: Distinguished from greggii , the only other small species of Pseudopsallus with a strongly produced anteocular portion of the head, by the black antennal segment 1, small dark fuscous spots on the dorsoapical surface of the femora, smaller fuscous labium, more narrow vertex, and shorter anteocular portion of the head (measurement comparison in preceding species treatment).

DESCRIPTION: Male: Small, parallel­sided. MEASUREMENTS: Total length 3.38 (2.90 –3.85), length apex clypeus­cuneal fracture 2.31 (2.10–2.60), maximum width across hemelytra 1.20 (1.10 –1.34); head width 0.65 (0.59–0.69), vertex width 0.32 (0.30–0.33), anteocular length 0.28 (0.26– 0.30), eye width 0.15 (0.13–0.18), eye height 0.31 (0.28–0.33); antennal length segment 1: 0.22 (0.21–0.23), 2: 0.99 (0.86–1.14), 3: 0.72 (0.63–0.83), 4: 0.28 (0.24–0.35); labium length 1.68 (1.50–1.85), reaching middle of genital segment; pronotum width 1.00 (0.86– 1.08), pronotum length 0.46 (0.41–0.50). COLORATION (fig. 1): yellowish white; head with variable dark fuscous markings anterior of antennal fossa, four pairs of marks laterally on frons and temporal areas, collum entirely dark; antenna dark fuscous to black; labial segments 3 and 4 fuscous; pronotum with black anterior margin, dark fuscous calli, brownish gray infuscation on propleuron and disk; mesoscutum and scutellum with large, paired bilateral dark fuscous marks; hemelytra brownish gray infuscation with pale yellowish white sutures and veins; cuneus dark fuscous with pale apex; membrane faintly infuscate, veins pale yellowish white; abdomen dark fuscous with variable yellowish brown mottling; tumid setose region on left side of genital aperture; legs with dark brown coxae; femora and tibiae yellowish brown with small scattered dark fuscous spots; tarsi dark fuscous. SURFACE AND VESTITURE: Subshining, smooth, dorsal vestiture of mixed scattered, broad, silvery, scalelike setae (fig. 8C) and moderately densely distributed, pale, reclining simple setae; tibiae with pale reclining setae, dark brown spines, and parallel rows of minute black spicules. STRUCTURE: Head triangular in dorsal view, length from apex of clypeus to vertex 90% of head width across eyes; frons gently rounded; clypeus obvious in dorsal view; anteocular distance in lateral view equal to 43 diameter of antennal segment 1, anteocular distance 15% greater than distance posterior of antennal fossae in dorsal view; eye height in lateral view equal to 82% of head height; pronotum subtriangular in dorsal view; corial margins parallel. GEN­ ITALIA: Genital segment: aperture without tergal processes (figs. 8F, 9A); left surface of segment, between midline and paramere insertion, tumid with stouter setae then on remainder of genital segment (figs. 8F, 9A). Left paramere: DLL small, length shorter than one­half width of paramere, apex somewhat pointed (figs. 8F, 9E); LAL broad basally, slightly attenuate, apex rounded (figs. 8F, 9F). Right paramere: MIF margin smooth, with two spinules distally, troughlike region ventral to MIF smooth (fig. 9C); DLR produced dorsal to MIF, DP slightly produced with several apical spinules (figs. 8F, 9B). Phallotheca: cylindrical with ovate aperture (fig. 9D). Vesica: apparently without spicules; ductus with aperture directed ventrally (fig. 9D).

Female: Macropterous. Similar to male in color, vestiture, and structure, except for smaller eyes, wider vertex, and slightly shorter hemelytra. MEASUREMENTS: Total length 2.97 (2.55–3.33), length apex clypeus­cuneal fracture 2.21 (1.89–2.45), maximum width across hemelytra 1.20 (1.02– 1.30); head width 0.68 (0.62–0.73), vertex width 0.36 (0.33–0.40), anteocular length 0.33 (0.29–0.36), eye width 0.15 (0.15– 0.17), eye height 0.31 (0.29–0.34); antennal length segment 1: 0.21 (0.19–0.23), 2: 0.89 (0.75–1.01), 3: 0.67 (0.56–0.78), 4: 0.26 (0.23–0.28); labium length 1.81 (1.66–1.95), reaching eighth abdominal sternite; pronotum width 1.00 (0.84–1.09), pronotum length 0.46 (0.40–0.53).

ETYMOLOGY: Named for its occurrence on species of Tiquilia Persoon (Boraginaceae) or coldenia and crinklemat.

HOST: T. palmeri (A. Gray) A. T. Richardson , Palmer’s crinklemat.

DISTRIBUTION: The Mojave desert in southeastern California east to western Arizona.

PARATYPES: USA: Arizona: La Paz Co.: Budweiser Spring, Kofa Mountains, March 20, 1980, J. T. and D. A. Polhemus, 15?, 12/ (JTP). California: Imperial Co. : Glamis, 2 mi W of, July 25, 1960, R. C. Dickson [?, handwritten], ex Coldenia palmeri , 1/ (UCR); Mt. Signal, at base of, 12 mi W of Calexico, March 30, 1974, J. D. Pinto, ex Coldenia palmeri , 5?, 5/ (UCR); same label data as holotype, 6?, 25/ [fig. 1 adult dorsal habitus photographs] (AMNH, USNM); Plaster City, 2.5 mi N of, May 1, 1952, Timberlake, ex Coldenia palmeri , 8?, 8/ (UCR); Truckhaven, April 15, 1949, Timberlake, ex Coldenia palmeri , 2?, 1/ (UCR). Inyo Co. : Wyman Canyon, White Mtns, July 9, 1976, S. and S. Frommer, ex light, 1? (UCR). Riverside Co.: Box Canyon, April 27, 1952, Timberlake, ex Coldenia palmeri , 1? (UCR). Mecca, ex Coldenia palmeri (UCR) : 3 mi E of, April 27, 1952, Timberlake, 11?, 3/; Box Canyon Wash, 5 mi E of, June 6, 1979, J. D. Pinto, 4?, 8/; 5 mi E of, October 5, 1976, J. D. Pinto, 4?, 7/. Snow Creek, White Water, 1500 ft, April 6, 1955, W. R. Richards, 1/ (CNC); Thousand Palms, April 3, 1955, W. R. Richards, 5?, 19/ (CNC).

Figures

Fig. 1. Habitus views of Calidroides, Chlamyopsallus, Pseudopsallus, and Strophopoda spp.

Fig. 6. Scanning electron micrographs of Pseudopsallus greggii?, Texas: Big Bend National Park A. Head and pronotum, dorsal view. B. Head, lateral view. C. Setae of corium, dorsal view. D. Pretarsus, lateral view. E. Mesothoracic spiracle and metathoracic scent efferent system, lateral view. F. Genital segment, caudal view. Scales as indicated.

Fig. 8. Scanning electron micrographs of Pseudopsallus tiquiliae?/, California: Mt Signal. A. Head and pronotum, dorsal view. B. Head, lateral view. C. Setae of corium, dorsal view. D. Pretarsus, apical view. E. Mesothoracic spiracle and metathoracic scent efferent system, lateral view. F. Genital segment, caudal view. Scales as indicated.

Fig. 9. Male genitalia of Pseudopsallus tiquiliae, California: Truckhaven. A. Genital segment, dorsal view. B–C. Right paramere. B. Medial view. C. Dorsal view. D. Vesica and phallotheca, dorsal view. E–F. Left paramere. E. Lateral view. F. Ventral view. DLL, dorsal lobe of left paramere; DLR, dorsal lobe of right paramere; DP, distal portion of dorsal lobe of right paramere; LAL, lateral lobe of left paramere; MIF, medial interior flange of right paramere. Scale 5 0.10 mm.