Tuxedo, SCHUH, 2001

SCHUH, RANDALL T., 2004, Revision of Tuxedo Schuh (Hemiptera: Miridae: Phylinae), American Museum Novitates 3435 (1), pp. 1-26 : 1-9

TUXEDO SCHUH

Tuxedo Schuh, 2001: 251 (n.gen.).

TYPE SPECIES: Chlamydatus bicinctus Van Duzee, 1914 .

DIAGNOSIS: Males recognized by the mostly castaneous, more rarely brownish, background coloration of the dorsum with the basal third of the cuneus always contrasting

1 George Willett Curator and Chair, Division of Invertebrate Zoology, American Museum of Natural History.

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Copyright © American Museum of Natural History 2004

ISSN 0003­0082

TABLE 1 Measurements of Tuxedo Species

TABLE 1 (Continued)

creamy white and frequently with a more or less complete creamy­white transverse fascia on the hemelytra near apex of scutellum (fig. 1). Sexual dimorphism moderate to very pronounced, male with elongate to long hemelytra, parallel­sided; females ovate (fig. 1). Antennal segment 2 sexually dimorphic, cylindrical and slightly enlarged in males, more slender and tapered toward base in females. Vesica in male (fig. 2) more or less J­shaped, very weakly twisted, with one or two variously ornamented spines apically; secondary gonopore close to apex.

Tuxedo spp. are most similar in type of sexual dimorphism (including antennae) to Coniferocoris Schwartz and Schuh (1999) and Pinophylus Schwartz and Schuh (1999) . The pattern of coloration in Tuxedo is most similar to that of Psallovius Henry (1999) and Ranzovius Distant (1893) . The sexual dimorphism (including antennae) in Ranzovius is unique and unlike any of the genera here compared with Tuxedo . The structure of the male genitalia in Tuxedo is closest to that of Coniferocoris spp. , but is also similar to that of Pinophylus spp. , Psallovius spp , and Ranzovius spp. Whereas Coniferocoris spp. and Pinophylus spp. are restricted to the Pinaceae , and Ranzovius spp. are commensal in spider webs, all known species of Tuxedo breed on angiosperms.

REDESCRIPTION: Male: Macropterous, of small to relatively large size, elongate, nearly parallel­sided; range total length 2.69–3.74, range length apex clypeus–cuneal fracture 1.84–2.39. COLORATION (fig. 1): Dorsum reddish or castaneous, with distinctive contrasting white or cream­colored maculae on cuneus and frequently on corium and cuneus, the transverse corial fascia just posterior to apex of scutellum. SURFACE AND VES­ TITURE (figs. 1, 3G, H, 4F, 5F): Dorsal body surface smooth, impunctate, polished, weakly to moderately shining. Dorsal vestiture of recumbent, simple, often shining setae; distal portion of dorsal surface of hind femur lacking row of spinules (fig. 5G). STRUCTURE: Head short, closely conforming to anterior margin of pronotum; posterior margin of vertex weakly elevated and round­ ed (figs. 3C, 4C, 5C); frons barely protruding beyond anterior margin of eyes; eyes large, head only slightly projecting below their ventral margin; antennae inserted just above ventral b margin of eyes (figs. 3A, 4A, 5A); antennal segment 2 cylindrical, not tapered, about the same diameter as antennal segment 1; labium reaching to about apex of hind coxae; claws elongate, smoothly curving, pulvilli small, located near base of claws, parempodia relatively short, setiform (figs. 3E, 4E, 5E); mesothoracic spiracle and metathoracic scent­efferent system as in figures 3D, 4D, and 5D; femoral trichobothria as in figure 4H; abdomen slender, genital capsule relatively small; sometimes ventral and posterior surface of genital capsule with a field of short, thickly set, peglike setae (fig. 5H). GENITALIA (fig. 2): Vesica more or less Jshaped, very weakly twisted, apically with one or two short spines; secondary gonopore very close to apex of vesica; phallotheca with apical portion elongate and slender, at nearly right angle to body of structure (figs. 3F, 4G); left paramere boat­shaped; right paramere lanceolate, with a nipplelike apex.

Female: Small to moderately robust, elongate ovoid; range total length 2.40–3.48, range length apex clypeus–cuneal fracture 1.67–2.43. COLORATION (fig. 1): Pattern of coloration often similar to male, but frequently showing sexual dimorphism, especially in coloration of antennal segments 1 and 2. SURFACE AND VESTITURE: As in male. STRUCTURE: Body form more compact than in male; submacropterous, hemelytra just covering abdomen; eyes smaller than in male, frons more prominently bulging anterior to eyes, head projecting below eyes by about one­third height of eye (figs. 3B, 4B, 5B); antennal segment 2 more slen­ der than in male, tapering toward base.

HOSTS: Most Tuxedo spp. breed on Ceanothus spp. (Rhamnaceae) , Cercocarpus spp. (Rosaceae) , or Quercus spp. (Fagaceae) , with a single species known from Fremontodendron spp. (Sterculiaceae) . All other recorded hosts occur with much lower frequency; some certainly do not represent breeding records.

DISTRIBUTION: Southern British Columbia, Canada , in the north, south to northern Baja California, Mexico , and east to western Colorado and New Mexico .

DISCUSSION: Schuh (2001) noted the similarity in coloration of Tuxedo spp. with species of Sejanus Distant from the Indo­West Pacific (see Schuh, 1984), suggesting by way of implication a possible placement it the Leucophoropterini . As mentioned in the diagnosis, the form of the male genitalia in Tuxedo is very similar to that of species placed in Coniferocoris and also has many similarities with that of Psallovius and Ranzovius . The type of sexual dimorphism is very similar to that of Coniferocoris and Pinophylus , although the genitalia of Pinophylus are not so similar to those of Tuxedo as are those of Coniferocoris , Psallovius , and Ranzovius .

Henry (1999) compared Psallovius and Ranzovius , treated the two as sister groups, and provided a phylogenetic analysis. Coniferocoris , Pinophylus , and Tuxedo were not documented at that time, but it would appear that the five genera may well form a monophyletic group based on the pattern of coloration and the form of the male genitalia. Henry (1999) rooted his tree with what he referred to as a ‘‘ Plagiognathus like’’ ancestor. Henry (1999) assumed that Psallovius and Ranzovius were members of the Phylini . The issue of whether they are members of the Phylini or the Leucophoropterini (as implied by Schuh) is a subject that could benefit from a more broad­based analysis, something that I do not propose undertaking at the present time.

All diagnoses in the present paper are written as applying only to male specimens. A key to the males is provided. The process of identifying females is best conducted through association with males. The reasons are that whereas the males of three of the known species have dark­colored antennae, all species but cruralis have pale antennae in the females, offering much less variation with which to discriminate the taxa. The coloration of the femora will assist in the descrimination of females of some species.

KEY TO MALES OF TUXEDO

1. Antennal segments 1 and 2 black, or nearly so; hemelytra with or without transverse fascia........................... 2

— Antennal segment 2 pale, segment 1 also usually pale; if segments 1 and 2 not entirely pale then never black; hemelytra always with a partial to complete pale, transverse fascia at level of apex of scutellum... 4

2. Clavus uniformly dark, never with a pale marking at level of apex of scutellum, or if with a faint marking then all femora and tibiae distinctly infuscate............ 3

— Clavus with at least a faint pale area at level of apex of scutellum, this area often heavily contrasting with dark background coloration of hemelytra; fore­ and middle femora and tibiae always pale; Oregon to Arizona; breeds on Quercus spp susansolomonae , new species

3. At least basal area of corium of paler coloration than remainder, sometimes much of corium pale, but hemelytra never with a pale, median, transverse fascia; all femora entirely infuscate; small to medium­sized species; widely distributed from British Columbia south to Baja California and east to Colorado and New Mexico; breeds primarily on Ceanothus spp. and Cercocarpus spp. ......... cruralis (Van Duzee)

— Corium and clavus uniformly dark, clavus occasionally with a limited pale area near apex of scutellum; femora infuscate only on distal half; large, heavy bodied species; central California, eastern Washington; breeds on Quercus spp. ............................... flavicollis (Knight)

4. Elongate slender species, ratio of length to width 4.2:1; northern California to Arizona; on Fagaceae elongatus , new species

— Not so elongate, ratio of length to width nev­ er more than 3.30:1............... 5

5. Coloration of dorsum generally uniformly castaneous or reddish brown except for cream­colored base of cuneus and pale transverse fascia at apex of scutellum 6

— Coloration of dorsum variably brown to pale, with weakly to moderately contrasting markings at base of cuneus and as transverse fascia at apex of scutellum; Arizona, southern California; breeds on Quercus spp. ................ nicholi (Knight)

6. Hind femur entirely castaneous; smaller species, length 2.76–2.92, width pronotum 0.79–0.91; transverse fascia at apex of scutellum only weakly developed on corium; vesica in male as in figure 2; California, Oregon, Nevada; breeds on Ceanothus spp. and Cercocarpus spp. ......................... bicinctus (Van Duzee)

— Hind femur castaneous on distal one­half, pale proximally; larger species, length 3.01–3.22, width pronotum 0.98–1.06; transverse fascia at apex of scutellum fully developed on clavus and corium; vesica in male as in figure 2; southern California; breeds on Fremontodendron californicum ............................... drakei , new species

Tuxedo bicinctus (Van Duzee)

Figures 1–3

Chlamydatus bicinctus Van Duzee, 1914: 30 (n.sp.).

Microphylellus minor Knight, 1929: 42 (n.sp.). NEW SYNONYMY.

Microphylellus bicintus: Pinto, 1982: 102–109 (host, phenology).

Tuxedo bicinctus: Schuh, 2001: 252 (n.comb.).

LECTOTYPE: Chlamydatus bicinctus Van Duzee (here designated): Male: [ USA:] ‘‘ San Diego, Calif., V­6–1913, EP Van Duzee’ ’. Deposited in the California Academy of Sciences.

HOLOTYPE: Microphylellus minor Knight : Male: ‘‘Fresno, Calif., June 20, 1926, C. J. Drake. ’’ Deposited in the National Museum of Natural History , Washington, D.C.

DIAGNOSIS: Recognized by the very small size in combination with the pale antennae, the cream­colored fascia across the hemelytra somewhat weakly developed on the corium, the entirely castaneous hind femora, and the habit of breeding on Ceanothus spp. and Cercocarpus spp. Most similar in appearance to T. drakei and T. susansolomonae , but distinguished from drakei by the hind femur in that species being castaneous only on the distal half, and from both by their slightly to significantly larger size, as well as different host preferences. Vesica in male also distinctive, with a single, short, bifid apical spine very similar to that found in drakei , but readily distinguished from that species through comparison of the overall conformation of the vesica.

REDESCRIPTION: Male: Small; total length 2.76–2.92, length apex clypeus–cuneal fracture 1.84–2.02, width pronotum 0.79–0.91. COLORATION (fig. 1): Background coloration castaneous; basal one­third to one­half of cuneus creamy white; clavus always with a rounded pale area just posterior to apex of scutellum, sometimes more extensive and nearly parallel­sided and extending onto adjacent corium and forming a more­or­less continuous, pale, transverse fascia reaching to the R­M vein; antennal segments 1 and 2 usually entirely pale, segments 3 and 4 appearing darker; labial segments 2 and 3 pale; all coxae, fore­ and middle femora, and all tibiae pale; hind femora entirely castaneous. SUR­ FACE AND VESTITURE (fig. 1, 3G, H): Pronotum moderately shining, remainder of dorsum weakly shining. STRUCTURE: Body appearing somewhat flattened; head as in figure 3A, C; meso­ metathoracic pleuron as figure 3D; claws as figure 3E. GENITALIA (figs. 2, 3F): Body of vesica relatively stout in comparison to most other Tuxedo spp. , most similar to Tuxedo drakei , but shorter and with basal region more strongly curving than in that species; vesica with a single, bifid apical spine extending only short distance beyond secondary gonopore and envelope of vesica, of form very similar to that of T. drakei ; apical vesical spine in T. cruralis also of similar appearance, but longer and curving, and vesica not so stout as in that species.

Female: Small; total length 2.40–2.62, length apex clypeus–cuneal fracture 1.67– 1.82, width pronotum 0.82–0.90. COLOR­ ATION (fig. 1): As in male, except transverse fascia at apex of scutellum always broad and intense on clavus and most frequently complete and extending to and including costal vein. SURFACE AND VES­ TITURE: As in male. STRUCTURE (fig. 3B): As in generic description.

HOSTS: Ceanothus spp. (Rhamnaceae) and Cercocarpus spp. (Rosaceae) . Also recorded from Phoradendron spp. , although this may not be a breeding host.

DISTRIBUTION: Central Oregon and southern Idaho south to San Diego County, California , and east to southern Utah.

DISCUSSION: Van Duzee (1914) described Chlamydatus bicinctus from ‘‘numerous specimens taken on Ceanothus in the spring, March to June’’. I have examined 4 male and 15 female specimens collected by Van Duzee; one male is labeled lectotype, one female as allotype, the remainder being labeled as paratypes. The ‘‘type’’ labels were apparently placed on the specimens by Van Duzee subsequent to the publication of the description of the taxon, but my research suggests that no lectotype designation was ever published. I am therefore formally designating the male specimen from the California Academy of Sciences bearing a red lectotype label as the actual lectotype for this species .

Knight (1929) indicated that C. J. Drake had collected the nominal species bicinctus Van Duzee and minor Knight on different hosts at the same locality at Fresno, California. According to Knight, his species minor was smaller, with a more porrect head, and with the tylus projecting distinctly forward, and with the vertex also somewhat wider than in bincinctus. It is my view that Knight was certainly correct that Drake had collect­ ed two species at Fresno on June 20, 1926. Knight, however, did not make it clear how he fixed the identity of bicinctus . Neither did he say how it was that he knew that Drake had collected what he thought were different species of bugs on different plants. The specimen labels contain no such information and there is no chance that two species of this genus could be distinquished as different in the field except by the fact that they occupied different hosts. I have compared the Van Duzee material of T. bicinctus mentioned above with the holotype and allotype of minor and conclude that the species Knight identified as bicinctus is actually undescribed, and that his species minor is actually bicinctus , and therefore a junior synonym. The identity of bicinctus sensu Knight is discussed below un­ der Tuxedo drakei , new species.

Pinto (1982) described the phenology of T. bicinctus on Ceanothus crassifolius at a site in Riverside County in southern California.

SPECIMENS EXAMINED: USA: California: Alameda Co.: Oakland, June 1, 1935, E. S. Ross, 13, 2♀ (CAS). Butte Co.: Oroville, May 1, 1925, H. H. Keifer, Ceanothus cuneatus (Rhamnaceae) , 53, 2♀ (CAFA, CAS). Fresno Co.: Fresno, June 20, 1926, C. J. Drake, 1♀ (CNC). Fresno, June 29, 1926, C. J. Drake, 13, 2♀ (USNM). Humboldt Co.: Beatrice, June 30, 1940, B. P. Bliven, 23 (CAS). Blocksburg, June 24, 1951, B. P. Bliven, 13 (CAS). Bridgeville, June 20, 1959, Kelton and Madge, 13 (CNC). Carlotta, July 23, 1967, 13 (CAS). McCann, July 19, 1959, Kelton and Madge, 13 (CNC). Redcrest, July 8, 1973, B. P. Bliven, 13 (CAS). Kern Co.: 20 km W of Wofford Heights on Rt 155, 1500 m, July 26, 1999, M. D. Schwartz, Cercocarpus betuloides (Rosaceae) , 3♀ (CNC). 7 mi W of Wofford Heights on Rt 155, 1520 m, June 26, 1999, M. D. Schwartz, Arctostaphylos sp. (Ericaceae) , 1♀ (CNC). near Kernville, Headquarters Camp, Sequoia National Forest, June 18, 1993, W. F. Chamberlin, 1♀ (TAMU). Lassen Co.: 9 mi W of McArthur, 1280 m, July 6, 1979, R. T. and Joe Schuh, Cercocarpus betuloides (Rosaceae) , 23, 2♀ (AMNH). Blue Lake, July 19, 1947, R. L. Usinger, 1♀ (UCB). Los Angeles Co.: Claremont, C. F. Baker, 13 (HELSINKI). Flintridge, May 10, 1955, C. L. Hogue, 13 (LACM). Los Angeles, April 1, 1900, Koebele, 1♀ (CAS). Pasadena, June 5, 1909, Grinnell, 1♀ (CAS). Marin Co.: Mt. Tamalpais, June 28, 1918, Mariposa Co. : Yosemite National Park, Glacier Point, 7214 ft, July 3, 1946, H. P. Chandler, 13 (CAS). Medera Co. : Coarsegold, June 29, 1946, R. L. Usinger, Phoradendron sp.(Loranthaceae) , 5♀ (UCB). Mendocino Co.: Eel River R.S., Mendocino National Forest, June 12, 1972, J. Doyen, Cercocarpus betuloides (Rosaceae) , 7♀ (UCB). Modoc Co.: Fandango Pass Summit, Warner Mts., 1890 m, July 3, 1979, R. T. Schuh and B. M. Massie, Cercocarpus ledifolius (Rosaceae) , 103, 17♀ (AMNH). Monterey Co.: Bryson, April 14, 1917, E. P. Van Duzee, 1♀ (CAS). Napa Co.: Soda Creek, May 3, 1931, H. H. Keifer, Cercocarpus sp. (Rosaceae) , 43 (CAFA). Orange Co.: 1.5 mi E of San Juan Campground, Cleveland Natl. Forest, May 12, 1978, J. D. Pinto and R. T. Schuh, 13 (AMNH). Riverside Co.: Menifee Valley (hills on W end), 1800 ft, April 29, 1979 – May 22, 1979, J. D. Pinto, Ceanothus crassifolius (Rhamnaceae) , 293, 32♀ (UCR). Pinon Flat, San Jacinto Mts., May 21, 1940, R. L. Usinger, Phoradendron juniperinum (Loranthaceae) , 13, 4♀ (UCB). San Jacinto Mountains, jct of Poppet Flat Rd and Rt 243, 1270 m, May 20, 2000, M. D. Schwartz, Cercocarpus sp. (Rosaceae) , 43 (CNC). SE of Murietta, road to Tenaja Fire Station, 1500 ft, May 13, 1978, J. D. Pinto, 1♀ (UCR). Tenaja Road W of Murietta, 410 m, May 12, 1978, J. D. Pinto and R. T. Schuh, Ceanothus crassifolia (Rhamnaceae) , 153, 14♀ (AMNH). San Bernardino Co.: Cajon Pass, jct Rts I­15 and 138, 1030 m, May 2, 1985, R. T. Schuh and B. M. Massie, Cercocarpus betuloides (Rosaceae) , 73 (AMNH). Lytle Creek, June 8, 1928, E. P. Van Duzee, 13 (CAS). San Diego Co.: Laguna Mts., Kitchen Creek Rd 1.4 mi N of Rt 8, 1100 m, May 21, 2000, M. D. Schwartz, Ceanothus sp. (Rhamnaceae) , 13, 2♀ (CNC). No specific locality, March 29, 1914 – June 1, 1914, E. P. Van Duzee, paratypes: 13, 11♀ (CAS). San Diego, May 17, 1913, W. S. Wright, 13, 3♀ (CAS). Tecate Peak, 1885 ft, June 2, 1980, Brown and Faulkner, 23 (SDNH). San Luis Obispo Co.: 10.5 mi SE of Santa Margarita , May 16, 1980, J. D. Pinto, 1♀ (UCR). Arroyo Grande Creek SW of San Luis Obispo, 160 m, May 8, 1985, R. T. Schuh and B. M. Massie, Ceanothus cuneatus (Rhamnaceae) , 23 (AMNH). San Mateo Co.: Lake Pilarcitos, June 25, 1966, C. W. O’Brien, 73, 7♀ (UCB). Santa Barbara Co. : Santa Cruz Island, Can del Medio, April 28, 1969, D. S. Horning, 13 (UCD). Upper Oso Campground off Rt 154, 310 m, May 7, 1985, R. T. Schuh and B. M. Massie, Cercocarpus betuloides (Rosaceae) , 23, 10♀ (AMNH). Shasta Co.: 1 mi E of Montgomery Creek, May 27, 1968, R. M. Brown, 1♀ (CAS). 7.6 mi N of Manton, 1138 m, July 10, 1980, R. T. Schuh and G. M. Stonedahl, Cercocarpus alnifolius (Rosaceae) , 2♀ (AMNH). Hat Creek P.O., June 20, 1955, J. W. McSwain, 13, 6♀ (UCB). Paynes Creek, June 18, 1959, Kelton and Madge, 9♀ (CNC). Siskiyou Co.: 2 mi SE of Lava Beds Natl. Monument, 5000 ft, June 26, 1979, M. D. Schwartz, Cercocarpus ledifolius (Rosaceae) , 33, 5♀ (OSU). Lava Beds Natl. Monument near Headquarters, 1560 m, June 26, 1979, R. T. and Joe Schuh, Cercocarpus ledifolius (Rosaceae) , 23, 10♀ (AMNH). McCloud, July 23, 1918, E. P. Van Duzee, 1♀ (CAS). Medicine Lake Road, 4800 ft, June 26, 1979, G. M. Stonedahl, Cercocarpus ledifolius (Rosaceae) , 9♀ (OSU). Sisson, July 25, 1918, E. P. Van Duzee, 13, 4♀ (CAS). Idaho: Owyhee Co.: Silver City, 6200 ft, July 8, 1973, P. W. Oman, 13 (OSU). Nevada: Clark Co.: Charleston Peak, July 21, 1982, J. T. Polhemus, Cercocarpus sp. (Rosaceae) , 23, 7♀ (JTP). Oregon: Grant Co.: Malheur National Forest, T14S R33E Sec 15, July 21, 1979, M. D. Schwartz, Cercocarpus ledifolius (Rosaceae) , 53, 8♀ (AMNH). Harney Co.: 11 mi E of Frenchglen, 6000 ft, July 16, 1957, J. D. Lattin, Cercocarpus sp. (Rosaceae) , 43, 18♀ (OSU). Klamath Co.: 28 mi SE of jct Rts 97 and 31, 4910 ft, July 25, 1979, M. D. Schwartz and G. M. Stonedahl, Cercocarpus ledifolius (Rosaceae) , 63, 3♀ (AMNH). 28 mi SE of jct Rts 97 on Hwy 31, 4910 ft, July 25, 1979, M. D. Schwartz, G. M. Stonedahl, Cercocarpus ledifolius (Rosaceae) , 63, 5♀ (OSU). Base of Bly Mountain, June 25, 1961, Joe Schuh, Cercocarpus ledifolius (Rosaceae) , 1♀ (AMNH). Lake Co.: 24 mi SE of LaPine, T24S R13E Sec. 24, July 12, 1957, G. F. Kraft, Cercocarpus ledifolius (Rosaceae) , 1♀ (OSU). Wheeler Co.: 4.5 mi S of Mitchell on Summit Prairie Road, June 22, 1979, R. T. Schuh, Cercocarpus ledifolius (Rosaceae) , 1♀ (AMNH). 4.5 mi S of Mitchell on Summit Prairie Road, June 27, 1979, M. D. Schwartz, Cercocarpus ledifolius (Rosaceae) , 23 (OSU). Utah: Grand Co.: 11 mi SE of Rt 163 toward Dead Horse Point on Rt 313, 5200 ft, June 11, 1982, M. D. Schwartz, Cercocarpus ledifolius (Rosaceae) , 13, 1♀ (AMNH).

Figures

Fig. 1. Habitus views of Tuxedo spp. See appendix 1 for localities.

Fig. 2. Male genitalia of Tuxedo spp. See appendix 1 for localities.

Fig. 3. Scanning electron micrographs of T. bicinctus from USA: Oregon: Harney Co.: Malhuer National Forest (3) and USA: Oregon: Harney Co.: 11 mi E of French Glen (♀). A. Head, 3, lateral view. B. Head, ♀, lateral view. C. Head, 3, anterior view. D. Mesothoracic spiracle and metaepisternal scent efferent system, lateral view. E. Pretarsus, frontal view. F. Genital segment, dorsal view of left paramere, phallotheca, and right paramere. G. Setae on hemelytra. H. Setae on hemelytra, detail of microstructure. Scales as indicated.