I $ S. Grozeva &N. Simov (Eds) 2008 ADVANCES IN HETEROPTERA RESEARCH Festschrift in Honour of SO''' Anniversary ofAlkhailJosifov, pp. 339-348. © Pensoft Publishers Sofia-Moscow

The first Chorosoma (: : Rhopalinae: Chorosomatini) from the New World: Chorosoma josifovi nov. sp.

M.D. Schwartz1, C.W. Schaefer2 &J.D. Lattin3

'Agriculture & Agri-Food Canada, Research Centre, 960 Carling Avenue, Ottawa ON K1A 0C6, Canada. E-mail: [email protected] 'Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs CT 06269-3043, U.S.A. E-mail: [email protected] 1 Department of Botany and Plant Pathology, Oregon State University, Corvallis OR 97331-2902, U.S.A.

ABSTRACT

Except for one Ethiopian species, Chorosoma until now has been an exclusively Palearctic genus. Fifty years ago, a new species was discovered in the northwestern United States; we name this species Chorosoma josifovi in honor of the 80th birthday of Michail Josifov, TOO described C. gracile among his many other contributions to our knowledge of Het- 'roptera. The new species resembles the widespread C. schillingii in some features, and resembles other species in some other features.

INTRODUCTION

I Tn ' mz six species in Chorosoma are all Palearctic (GOLLNER-SCHEIDING 1983; DOLL- NG 2006) except C. xenocles LINNAVUORI, which occurs in Ethiopia (LINNAVUORI "*• In 1956, several specimens of the genus were collected in the state of Idaho, , •"•A..; and since then others have been found in Oregon and Nevada, all adjacent coll"e, s in the far northwest of the United States (Fig. 1). So far, there have been no actions of the species from Utah or from north of Oregon—that is, from central ; Wash ngton state north through the dry land areas of British Columbia and the Yukon 340 M.D. Schwartz, C.W. Schaefer &.J.D. Latrin

I

Fig. 1: Map of the distribution of Chorosoma josifovi nov. sp. (Pacific Ocean at lower left)

Territory of western Canada into Alaska (HENRY 1988; SCUDDER 1997; MAW et al. 2000; PAIERO et al. 2003). Comparison of determined specimens, illustrations, and descriptions of the six described Chorosoma species with the North American specimens, makes clear that these collections represent a new species. We describe this new species here, and name it for Dr. Michailjosifov, in honor of his many fine contributions to the study of Het- eroptera (including the description of Chorosoma gracile) and in happy celebration of his 80th birthday. Because most other species of Chorosoma occur in the Palearctic, we assume that the ancestor of this new species has immigrated to the New World from the Old. .Although members of the genus occur widely throughout eastern Russia and the former eastern Soviet Republics, and occur also in Mongolia, no species of Chorosoma appears to have been recorded so far from far-northeastern Russia, near Beringia (see MARTYNOVA 1975; GOLLNER-SCHEIDING 1983; VINOKUROV 1988; DOLLING 2006), where the two continents were most recently connected during the Pleistocene, although VINOKUROV & KANYUKOVA (1995) record it from Siberia. Indeed, the preference of Chorosoma for very dry grasslands suggests a movement into the New World long ago, before the growtn of the boreal forests. We are now studying the systematic and biogeographic relation ships and history of this new species, and we shall discuss these elsewhere (SCHAEFE , SCHWARTZ & LATTIN, in preparation). fig. The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 341

TAXONOMY

Chorosoma josifovi nov. sp. (Figs 2-5)

Description. Male: Gracile, abdomen extending beyond membrane of hemelytron for distance subequal to length of membrane; total length 10.14-11.60 mm (Fig. 2). COL­ ORATION: Entirely pale yellowish brown to sandy, except for reddish brown speckling on head, antennal segments I and II, and dorsum and sublaterally on venter of abdomen; hemelytron and membrane hyaline, embolium and veins concolorous with rest of body, except sometimes faintly brown on radial vein; reddish brown lateral stripe between antennal insertion and eye and faint lateral stripe on abdominal venter submedial to connexivum; brown marks at base of suture between clypeus and mandibular plate, tip of labium, variable curved mark on medial periphery of callus, and short apical stripe dorsal to procoxal cleft and ventral to lateral carinate margin of pronotum; abdominal dorsum with prominent, wide, irregular, dark brown or black line just medial to con­ nexivum extending from tergum 1 (IT) (beside scutellum) to or slightly beyond 4T or ST, lines occasionally shorter or longer; some specimens with line anteriorly, between lateral fines; trichobothrial bases and abdominal spiracles not contrastingly colored; hind

Fig. 2: Chorosoma josifovi nov. sp., dorsal habitus. 342 M.D. Schwartz, C.W. Schaefer &J.D. Lattin

leg with dark brown on apicomedial surface of tibia, apical half of tarsomere I- tar III and pretarsus dark brown; tips of parameres dark brown to black. SURFACF VESTITURE: Discretely punctate, except cafiar region smooth and slightly n moderately densely covered with very short, appressed pale setae; setal bases of post 11 ' • pronotum often dark; antennal segment IV also with sparse, slightly longer, suberect apical portion of tibia, and tarsomeres with longer setae; medial surface of metatibia with erect, pale, bristlelike spines with length subequal to diameter of tibia. STRUCTURF' Lateral margins of head subparallel, except for bulge of eye; base of head slightly raised ridgelike, raised area extending forward sublaterally to enclose ocellar tubercle- tuber­ cle low. Eyes rather prominent, distance across them (lateral margin to lateral marrin) approximately 80% of maximum width of pronotum. Antenna relatively long, anten­ nal segment I thicker than others, subequal in length to head (clypeus to ocellar line) parallel-sided laterally, convex medially; II-IV slender, II slightly thicker proximally, IV slightly clavate, II=-III>I=IV (see Measurements). Labium reaching mesocoxa, segment I>II>IV>III. Pronotum narrow, broadened slightly caudally, lateral margins very sUghdv sinuate, posterior margin straight, callus broad, disc densely punctate, median carina faint, especially apicafiy; humeral area produced laterally into broad rounded tubercle. Scutel­ lum narrow apicafiy, slightly concave medially with faint median carina, apex rounded, Fig. 5: Ck distinctly concave, slightly raised. Apex of hemelytron membrane reaching apex of ab­ dominal segment V. Pretarsus with strongly curved claw, reaching beyond apex of broad and apicafiy attenuate pulvillus. GENITALIA: Pygophore with rather widely spaced lateral and paralateral lobes; both lobes longer than medial lobe (Figs 3, 4). Parameres distinctly narrow apicafiy, tip strongly sclerotized, curved laterally (Kg. 5). Aedeagus with tighdy coiled vesical base; apical region with sclerotized and membranous lobes. Female: Same as male, but larger (Table 1); apex of hemelytral membrane reaching middle of ST.

medial lobe paralat. lobe

lar. lobe

Fig. 3: Chorosoma josifovi nov. sp., pygophore, lateral view; scale = 0.10 mm. Fig. 4: Chorosoma josifovi nov. sp., pygophore, dorsal view; scale = 0.10 mm. The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 343

Fig. 5: Chorosoma josifovi nov. sp., right paramere, lateral view; scale = 0.10 mm.

TERMINALIA: Proctiger prominently exposed ventrally, covered dorsally by exposed 8T and 9T; ovipositor concealed by seventh sternum. MEASUREMENTS: Table 1. ABBREVIATIONS: CNC, Canadian National Collection, Ottawa; CWS, col­ lection of Carl W. Schaefer, Storrs; OSUC, Oregon State University, Corvallis; UIM, University of Idaho, Moscow; UNV, University of Nevada, Reno; USNM, United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. HOLOTYPE: $. [U.S.A.] 1) "Blow Sand Mtns./ Churchill Co.NV/T16N,R30E [39.204442, -18.591547,1317m]/ viii-2-1979.2) R. C. Bechtel/ R. W. Rust/ Collectors. 3) Oryzopsis hymenoides. 4 [red] HOLOTYPE/ Chorosoma josifovi/ Schwartz, Schaefer, & Lattin." Deposited in USNM. PARATYPES [22 specimens with blue PARATYPE labels; no specimen is in perfect condition]: U.S.A.: Idaho: Fremont Co.: Sand Dunes.Saint Anthony [43.962428, -111.854627,1488m], 16 July 1956, W. L. Barr, Id', 1? (CNC), 3-5 (UIM). Nevada: Churchill Co.: same labels as holotype, 1<$, 2$ (CWS), 12 (OSUC), 3d, 2$ (USNM); Blow Sand ML, 28 mi SSE Fallon, June 1979, R. W. Rust, 12 (OSUC, UNV), 22 (USNM). Oregon: Harney Co.: Alvord Desert Sand Dunes, 18 June-1 July 1986, N. Cobb, ex: Oryzopsis hymenoides, Sporobolus airoides, 1

Table 1. Measurements (mm.) of Chorosoma josifovi nov. sp. Specimens from Idaho (U.S A) d'ff somewhat from those from Oregon and Nevada, and are listed separately.

Male Female •• Idaho Oregon/Nevada Idaho Oregon/Nevada ' (N=4) (N=4) (N=l) (N=2) Total length 11.31 (10.75-11.60) 10.29 (10.14-10.53) 14.80 13.33 (12.74-ll9ir~ Length: head to 8.25 (7.80-8.80) 7.45 (7.28-7.54) (N=3) 9.70 8.84 (8.19-9.49) apex hemelytron Head length- 1.33 (1.25-1.40) 1.29 (1.29) (N=2) 1.50 1.53 (N=l) Ocular distance 1.17(1.15-1.18) 1.13 (1.09-1.16) (N=2) 1.33 1.23(1.19-1.32) (from outer edge of each eye) Interocular width 0.69 (0.66-0.70) 0.70 (0.66-0.73) (N=2) 0.83 0.89 (0.86-0.91) Antennal segments: I 1.52 (1.48-1.60) 1.41 (1.39-1.41) (N=2) 1.68 1.75 (N=l) n 2.32 (2.25-2.45) 2.05 (1.98-2.11) (N=2) 2.48 2.38 (N=l) in 2.17 (2.05-2.30) 1.90 (1.82-1.98) (N=2) 2.15 2.05 (N=l) IV 1.58 (N=l) 1.64 (1.52-1.75) (N=2) 1.45 (N=l) Pronotal length 1.38 (1.30-1.38) 1.19 (1.19) (N=2) 1.66 1.39 (1.32-1.45) Pronotal width 1.43 (1.41-1.45) 1.35 (1.32-1.39) (N=2) 1.70 1.64(1.52-1.75) Rostrum length 3.11 (3.00-3.20) 2.78 (2.54-2.94) (N=3) 3.10 2.96 (2.81-3.10) Scutellum length 0.81 (0.75-0.86) 0.71 (0.60-0.73) (N=3) 0.95 0.85 (0.79-0.92) (N=3) Scutellum width 0.49 (0.44-0.54) 0.56 (0.53-0.59) (N=3) 0.55 0.62 (0.61-0.63 (N=3)) "from tip of clypeus to anterior edge of ocellar line

1 spec, consisting of head and thorax only (CWS). Nevada: Churchill Co.: same labels as holotype, 4 spec, consisting of head and thorax only (all in CWS). VARIATION: There is some variation in coloring and patterning, most noticeably between the specimens from Idaho and those from Oregon and Nevada (see below). The Oregon specimens also show other variation. In particular, the two sublateral dark lines on the abdominal dorsum may extend only as far as the diird tergite (3T), or as far as TT.The latter Table 2. occurs in the male holotype (Oregon); the former in a female collected on the same day at the same place. In other specimens also collected with these (same place and time), die dark lines end at 4T or 5T; this occurs in the majority of specimens. When the dark lines extend the full length of the abdomen, the appears dark; nevertheless, the ground color is pale. Minor variation also occurs in the degree of overall paleness, and in the degree of darkness, and extent, of the various markings given in the Description. None or this variation is consistent by sex or population. The first Chorosoma (Hemiptera: lihopalidac: Rhopalinae: Chorosomarini) from the New World... 345

DIFFERENCES BETWEEN UNITED STATES POPULATIONS: There are some differences between the specimens collected in 1956 in Idaho (at 1488 m), and those collected later (1979,1986) in Oregon (at 1281 m) and Nevada (at 1317 m), although structurally they are the same. These populations are separated by habitat inhospitable to Chorosoma and are not close to one another (the closest localities are 322 km apart, and the farthest are 724 km apart).The specimens from Idaho are in general darker than the Oregon-Nevada specimens. They also differ with respect to several measurements (Table l).Tne Idaho specimens are somewhat, but consistently, larger. Perhaps of greater significance, would be differences in absolute and relative lengths of head and antennal segments.These appear to be important in distinguishing Chorosoma species (see key in LINNAVUORI 1976). We have calculated several ratios to aid discussion (Table 2). The head length/ antennal-segment I ratios do not differ gready, either by sex, United States state, or date of collection (the last two of course are a single variable). This ratio indicates that the first antennal segment is longer, but not greatly longer, than the head (measured from clypeal tip to an imaginary line drawn across anterior edge of ocelli). Similarly, the first antennal segment is about two-thirds the length of the second, and there is little variation among the North American specimens. None of these measurements (Table 1) or ratios (Table 2) suggests that these different North American populations are not conspecific.There may be significant interspecific dif­ ferences in the genus, however, which we shall consider in the future (in preparation). HABITAT AND FOOD PLANTS: AU specimens of Chorosoma josifovi weie col­ lected on dune grasses in or on sand dunes, from -1280-1490 m elevation.The recorded food plants are the native grasses Oryzopsis hymenoides (ROEMAR & SCHULTES) RICKER fricegrass") (now Achnatherum hymenoides (ROEMER AND SCHULTES) BARKWORM) and tyorobohts airoides (TORR.) TORR. ("finetop saltgrass"). Both grasses grow on sandy, or very sandy, soil (FERNALD 1950; GLEASON & CRONQUIST 1991); S. airoides in alkaline meadows (GLEASON & CRONQUIST 1991) and saline flats (FERNALD 1950). With respect to habitat and food plant, Chorosoma josifovi is similar to other species of Chorosoma, at least to the European ones. C. xenocles was collected in (near?) Addis Ababa, Ethiopia (LINNAVUORI 1976), whose elevation is, for the most part, less than 3000 m. COMPARISON WITH OTHER SPECIES: The following discussion is based on identified specimens of Chorosoma brevkolle HSIAO, C. graci/e JOSIFOV, C. longkolle REUTER, wd C. schillingii (SCHILLING); in addition, for these species and for C. xenocles LINNA-

lable2. Ratios of several measurements of United States Chorosoma josifovi nov. sp. (data from Table 1).

Idaho specimens Oregon ScNi ,vad t specimens Male Female Male Female "tad length/antennal 1 0.86 0.89 0.91 0.86 - Wnnall/II 0.66 0.68 0.69 0.74 lead to hemelytron apex/body length 0.73 0.66 0.72 0.66 346 M.D. Schwartz, C.W. Schaefer &.J.D. Lattin

VUORI, we have trusted descriptions, illustrations, and characters in LINNAVUORI (197^ Distinguishing among these species is not easy, because in all species there is considerahl intraspecific variation in color and in structure. For example, the most widespread snec' ' Chorosoma schillingii, varies in size: we have identified specimens from "England" (no furth data) that are considerably larger than other specimens from Kazakhstan (England mal 14.43 mm, Kazakhstan male 12.35; England female 17.42, Kazakhstan female 14.43) And judging from the illustrations of parameres and genital capsules in JOSIFOV (1968, Figs 7-9 13-17), LINNAVUORI (1976, Figs 12c, d), and PEHLIVAN (1981, Fig. 54), these structures also vary considerably. We shall discuss these differences, similarities, and intraspecific variations elsewhere (SCHAEFER, SCHWARTZ, & LATTIN, in preparation). Chorosoma josifovi differs from other Chorosoma species chiefly and most consis­ tently in being considerably paler (especially the Oregon-Nevada specimens) and more uniformly colored. In particular, the dorsum of the abdomen in the other species we have seen is darker than in C. josifovi. Because these other species occur in the same sort of dry habitat as C. josifovi, the paleness of the latter cannot be attributed entirely to its habitat; therefore, this color difference appears to be significant. Chorosoma josifovi is of moderate size (Table 1), not much different in length from the other species, except C. hrevkolle,wbich. is larger (17 mm; HSIAO 1964; the"l.'7" in the figure caption is a misprint); some C. schillingii are large, too: a female from "England" is 17.42 mm long, but other specimens are of average length (11-14 mm), like other species. Although except for color, distinguishing C. josifovi from other species is not easy, there are more subtle consistent differences, especially in genitalia, that convince us the New World specimens represent a new species. These differences too will be discussed elsewhere. In Chorosoma schillingii, the two sublateral dark lines on the abdomen's dorsum extend to the tip of the abdomen or onto the sixth tergite; in other species they usually end no more posteriorly than on the fourth or fifth tergite. However, the variation noted above renders this character less reliable than may have been thought. There are several keys to the species of Chorosoma: MARTYNOVA (1975), LINNA­ VUORI (1976), and PUTSHKOV (1986). Species are often separated in these keys not by distinct discrete differences but by relative differences such as antennal ratios. But here too confusion may occur: for example, antennal IV of C. hngkolk is "fast so lang wie III" (JOSIFOV 1968, p. 256); or equal in males but III shorter in females (MARTYNOVA 1975, PUTSHKOV 1986); or, the ratio of the two is 48: 58 (III: IV) (LINNAVUORI 1976, key, couplet 3). Also used to distinguish species is the relative length of the antenna, usually measured by the antennal I: head length ratio. In this regard the comparative and total lengths of the antennal segments of C. josifovi are very close to those of C. longtcolle (compare our Fig. 3 with JOSIFOV 1968, Fig. 1). The paramere of Chorosoma josifovi (Fig. 5) is longer and more slender than tna of C. longicolle, and bent laterally at the tip more than is the paramere of C. longicol e, whose paramere tip is bent almost dorsally, along the main plane of the paramere s bo • In these respects the C. josifovi paramere more closely resembles those of Cgracile, schillingii (JOSIFOV 1968, Figs 13-22; LINNAVUORI 1976, Figs 12c, d), and C. xenocles The first Chorosoma (Hemiptera: Rhopalidae: Rhopalinae: Chorosomatini) from the New World... 347

(LINNAVUORI 1976, Fig. 12b). In all species there is ventrafiy a median extension of the male's genital capsule (actually, the fused cuplike sclerite and median projection: see SCHAEFER 1977,1978). Like the paramere, this is more slender and longer in C. josifovi (Figs 3,4) than in C. longicolle, in the former (and in C. schillingii [JOSIFOV 1968, Fig. 7]) it is about three-fourths of the parameres length, but slightly more than half that length in C.gracile (JOSIFOV 1968, Fig. 10) and in C. longicolle. The paralateral lobe (Figs 3,4) is less prominent than in C. gracile (JOSIFOV 1968, Figs 10-12) and C. schillingii (JOSIFOV 1968, Figs 7-9; SCHAEFER 1978, Fig. 3). However.it is more prominent than that of C. longicolle (MARTYNOVA 1975, Fig. 9; CWS, personal observation). Finally, MARTYNOVA (1975) suggests that C. brevkolle is actually a composite of C. macilentum and one or more other described species. The close similarity of the species and the great intraspecific variation, point clearly to a need for a generic revision of Chorosoma, which must include an analysis of populations over the entire range of the genus. We shall discuss this need in greater detail elsewhere. Note: Part of this paper was presented at the Fourth European Hemiptera Congress (Ivrea, Italy, September 2007), by CWS.

ACKNOWLEDGEMENTS

We thank W L. Barr, University of Idaho (UIM) and R. W Rust, University of Nevada, Reno for originally bringing the specimens from Idaho and Nevada respectively to our attention. We also appreciate the assistance extended to us by R. G. Foottit for access to the collection and facilities of the Canadian National Collection. We are grateful also to V. Kask and J. O'Donnell (University of Connecticut) and B. B. Hall (Oregon State University) for the dorsal habitus illustrations; to Fedor Konstantinov, Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZISP), for translating several Russian-language papers; to D. Lightfoot and N. Cobb (while at OSU) for their collecting at the Alvord Desert site in Oregon; and to Dr. I. M. Kerzhner (ZISP) for gently revealing to us several major and minor errors in this paper. And we thank T.J. Henry (U.S. National Museum, Smithsonian Institution) for the loan of specimens. Finally, CWS is grateful to the sponsors and organizers (and especially to Dr. Peter Mazzoglio) of the Fourth European Hemiptera Congress, for support which allowed him to present part of this paper at the Congress.

PE3IOME

^0 elaeMnAJipu, HaiviepeHU npegu 50 eogumi 6 ce8epo3anagHama nacm Ha CAIH,, ce onucBa • M mynaBaHemo Ha Hetoroptera. HoBuam Bug Manogo6aBa no Hakou 6eAe3u uiupokopa3npocmpaHeHUfl C. schillingii. ToBa e Bmopua npeg- crr*a8ume.\ Ha poga usBtH zpaHUUume Ha ITaAeaplunuka. 348 M.D. Schwartz, C.W. Schaefer &J.D. Lattin

REFERENCES

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