Leucotaraxis piniperda Gaimari & Havill 2021, comb. nov.

Leucotaraxis piniperda (Malloch), comb. nov. (Figs 1D, 2E–H, 3C, 3H) Leucopis piniperda Malloch 1921: 351. United States. Illinois: Urbana (HT ♀, INHS). References – McAlpine 1965: 709 (catalog); Cole & Schlinger 1969: 376 (note); Tanasijtshuk 2002: 203 (redescription), figs 29–30 (head), figs 31–34 (♂ genitalia). Kohler et al. 2008: 494 (biology, as Leucopis atrifacies); Grubin et al. 2011: 1410 (biology); Havill et al. 2018 (biology, population genetics); Neidermeier et al. 2020 (biology, biological control); Dietschler et al. 2021 (biology, biological control). Diagnosis. The lunule, face and parafacial are either entirely silvery-grey pruinose (Fig. 2F, eastern form), entirely brown to black pruinose (Fig. 2H, western form), or the lunule and face are brown to black pruinose with the parafacial silvery-grey pruinose (Fig. 2G, mixed form). The postpronotum (Fig. 2E) has 1 strong postpronotal seta, and no additional setulae longer than those on the scutum. In the male (see figs 31–34 in Tanasijtshuk 2002), the epandrial complex is simple, small, and yellowish- to silvery-grey pruinose, with no central epandrial process; the surstylar lobe is short and stout; the phallus (Fig. 3C) is very long and thin in lateral view, more than 10 X longer than its height at the middle, and is gently curved throughout to a pointed tip. In the female (Fig. 3H), tergite 6 enlarged and about 1.5 X wider than long, tergite 7 absent (position represented only by tiny setulae), tergite 8 longer than wide, parallel-sided, with distal end enlarged; sternites as in Leucotaraxis atrifacies (Fig. 3F). Immatures. Egg and puparium as in Leucotaraxis atrifacies. Biology. Predators as larvae on species of Adelges and Pineus on Pinaceae (Abies, Picea, Pinus). Specific records (including our own rearings) include: Adelges piceae on Abies grandis; Adelges tsugae on Tsuga heterophylla (Kohler et al. 2008); Pineus boerneri on Pinus densiflora; Pineus coloradensis on Pinus sp. (a 5-needle pine, possibly Pinus lambertiana) and Picea engelmannii; Pineus pini on Pinus sylvestris; Pineus similis on Picea glauca; Pineus strobi on Pinus strobus (Tanasijtshuk 2002; Wantuch et al. 2019); Pineus sp. on Pinus sylvestris. They have also been collected as adults from Pinus contorta and Pinus resinosa. Remarks. Unfortunately, the few specimens from Oregon and Washington in the study by Kohler et al. (2008) were misidentified as Leucopis atrifacies. After examination of the specimens from this work and from Tanasijtshuk (2002), the color of pruinosity on the lunule, face and parafacial are variable, with a distinct distributional pattern. The specimens from eastern North America (Canada – New Brunswick, Quebec; USA – Connecticut, Illinois, Maryland, Minnesota, North Carolina, Ohio, Rhode Island, Virginia) all have uniformly silvery-grey pruinosity (Fig. 2F), while those in western North America (Canada – Alberta, British Columbia; USA – California, Colorado, Idaho, New Mexico, Oregon, Washington, Wyoming) all have uniformly brown to black pruinosity (Fig. 2H). The east-west divergence is also displayed in molecular data (see Havill et al. 2018). Only the specimens from Ontario, Canada, have a mixed state (Fig. 2G), with the lunule and face being brown to black pruinose with the parafacial silvery-grey pruinose. Both the male and female genitalia, as well as other external characteristics, make the identity of this species unambiguous despite this variation. The puparial stage is indistinguishable from that of Leucotaraxis atrifacies, both of which are only distinguishable from Leucotaraxis argenticollis by the shorter posterior spiracular tubercle that is not heavily sclerotized distally, and the slightly smaller size. Distribution. Canada (Alberta, British Columbia, New Brunswick, Ontario, Quebec, Saskatchewan). USA (Arizona, California, Colorado, Connecticut, Idaho, Illinois, Maryland, Massachusetts, Michigan, Minnesota, New Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Rhode Island, Utah, Virginia, Washington, West Virginia). Specimens examined. All specimens from Tanasijtshuk (2002) were examined, with additional specimens as follows. CANADA: ALBERTA. Seebe, 20.i.1971, ex. duff around C. com. infected P [inus]. contorta, #2670 (1 ♀, CNC). ONTARIO. Chalk River, 1.viii.1951, coll. J.M. Anderson, ex. Pissodes strobi (1 ♀, CNC). Ottawa, 29.vi.1994, coll. J.R. Vockeroth, ex. damp second-growther Acer - Betula woods (1 ♀, CNC). Sault Ste. Marie, 18. vii.1951, coll. L. Lyons, ex. red pine [= Pinus resinosa], No. R-70-19 (1 ♀, CNC). SASKATCHEWAN. Smeaton, 53.6181, –104.7409, 9.vi.2010, coll. N. Havill, ex. Pineus on Picea sp. (1 larva [molecular voucher], YPM; ENT764124). USA. CALIFORNIA. El Dorado Co., Eldorado National Forest, 38.9148, –120.3904, 31.v.2010, coll. Nathan Havill, Stephen Gaimari, ex. Pineus coloradensis on Pinus sp. (5-needle) (1 larva [molecular voucher], YPM; ENT790140). COLORADO. Boulder Co., Boulder, Flagstaff Canyon, 5800 ft., 10.vi.1961, coll. C.H. Mann, ex. on side of stream (1 ♂, CNC). Larimer Co., Zimmerman Lake trail, 40.5397, –105.8792, 27.vii.2016, coll. Nathan Havill, ex. Pineus coloradensis on Picea engelmannii (1 larva [molecular voucher], YPM; ENT857046). CONNECTICUT. Hartford Co., Hartford, Cedar Hill Cemetery, 41.7266, –72.6932, 5.vi.2016, coll. M.E. Montgomery, ex. Pineus strobi on Pinus strobus (8 larvae [molecular voucher], YPM; ENT961079–86). New Haven Co., Hamden, Exeter Rd. & Deerfield Dr., 41.426°N 72.936°W, 16.vi.2009, coll. M.E. Montgomery, ex. reared from Pineus sp. on Pinus sylvestris, N. Havill sample 09-134, Specimen #09E471 (1 ♂ [molecular voucher], CSCA). IDAHO. Idaho Co., Warm Springs Point, 13.viii.1969, 51-488b (2 ♀, USNM). ILLINOIS. Champaign Co. Urbana, 12.vi.1967, coll. J. Appleby, ex. larvae in wax mass of pine aphid (1 ♀ with puparium, 2 ♀, USNM). Morgan Co., Meredosia, 29.v.1917 (1 ♀, INHS). MARYLAND. Baltimore Co., Baltimore, Cylburn Arboretum, 39.532, –76.653, 28.v.2008, coll. Melody Keena, ex. Pineus boerneri on Pinus densiflora (7 larvae [molecular voucher], YPM; ENT856378–84). MASSACHUSETTS. Hampshire Co., Amherst, 42.3928, –72.5323, 20.v.2018, coll. Ryan Crandall, ex. Pineus strobi on Pinus strobus (1 larva [molecular voucher], YPM; ENT996097). Worcester Co., Worcester, 42.2790, –71.7733, 12.v.2018, coll. Felicia Andre, ex. Pineus strobi on Pinus strobus (1 larva [molecular voucher], YPM; ENT996096). MICHIGAN. Benzie Co., Lake Ann, 44.7436, –85.9131, 6.xi.2018, coll. Scott Lint, ex. Pineus strobi on Pinus strobus (2 puparia [molecular voucher], YPM; ENT996098–99). MINNESOTA. Beltrami Co., Bemidji, visitor center nr. Paul Bunyan statue, 47°28’13”N 94°52’43”W, 23.v.2009, coll. N. Havill, ex. old gall of Adelges sp. on Picea sp., N. Havill sample 09-123, Specimen #09E475 (1 ♂ [molecular voucher], CSCA). NEW MEXICO: San Miguel Co., Las Vegas Hot Springs, 7.viii, coll. H.S. Barber (1 ♂, USNM). NEW YORK. Cayuga Co., McIlroy Bird Sanctuary, 42.6661, –76.2834, 2.vii.2019, coll. Nicholas Dietschler, Marshall Lefebvre, Katharine O’Connor, ex. Pineus strobi on Pinus strobus (1 puparium [molecular voucher], CUIC). Monroe Co., Rochester, Highland Park, 43.1278, –77.6122, 8.vii.2017, coll. Cynthia Smith, ex. Pineus strobi on Pinus strobus (3 larvae, 13 puparia [molecular voucher], CUIC). Steuben Co., Addison, McCarthy Hill State Forest, 42.0975, –77.1938, 26.vii.2018, coll. Nicholas Dietschler, Marshall Bigler-Lefebvre, ex. Pineus strobi on Pinus strobus, (7 larvae, 2 puparia [molecular voucher], CUIC). Tompkins Co., Ithaca, Cornell campus, 42.4454, –76.4755, 3.vi.2016, coll. Nathan Havill, Mark Whitmore, Amalia Havill, ex. Pineus strobi on Pinus strobus (12 larvae [molecular voucher], YPM; ENT856925–36). Ulster Co., Woodstock, Overlook Mountain Forest, 42.0760, –74.1259, 31.v.2018, coll. Nicholas Dietschler, Marshall Bigler-Lefebvre, ex. Pineus strobi on Pinus strobus (5 larvae [molecular voucher], CUIC). NORTH CAROLINA. Macon Co., Highlands, Wightman College, 10. vii.1987, coll. W.W. Wirth, ex. UV light (1 ♀, USNM). Transylvania Co., Sapphire, 20.vii.1990, coll. W.W. Wirth, ex. UV light trap(2♀,USNM). OHIO. Morgan Co., Beverly, 39.6431, –81.7060, 26.vi.2019, coll. Page Weckbacher, ex. Pineus strobi on Pinus strobus (1 larva, 4 puparia [molecular voucher], YPM; ENT961088–92). Portage Co., Kent, 16.vii.1969, coll. T.P. Sluss, biological note 6914 (1 ♂, UAIC). OREGON. Benton Co., Corvallis, 12.v.1951, coll. R.G. Mitchell, ex. reared, predaceous on Adelges piceae (1 ♂, CNC). Multnomah Co., Portland, Grant Park, 45°32.450’N 122°37.800’W, 14.iv.2006, coll. G. Kohler, ex. reared from field collected puparium on Adelges tsugae (1 ♂ with puparium, CSCA), 12.v.2006, ex. flight caught (2 ♀, CSCA). PENNSYLVANIA. Greene Co., Mt. Morris, 39.7730, –80.1475, 19.vi.2015, coll. Andrew Liebhold, ex. Pineus strobi on Pinus strobus (2 larvae [molecular voucher], YPM; ENT856919, ENT856697). RHODE ISLAND. Washington Co., University of Rhode Island, Kingston, 12.vi.1982, coll. R. Casagrande (1 ♂, USNM). UTAH. Davis Co., Farmington Canyon, 40.9932, –111.8186, 5.ix.2017, coll. Danielle Malesky, ex. Adelges piceae on Abies lasiocarpa (1 larva [molecular voucher], YPM; ENT996095). VIRGINIA. Fairfax Co., Alexandria, 14.vi.1952, coll. W.W. Wirth (1 ♀, USNM), 15.vi.1952 (1 ♂, USNM). Montgomery Co., Blacksburg, Coal Mining Heritage Park, 37.1870, –80.427, 18.v.2010, coll. Melissa Fischer, ex. Pineus strobi on Pinus strobus. (9 larvae [molecular voucher], YPM; ENT856988–89, ENT856702–08). WASHINGTON. Clark Co., Vancouver, 45.7031, –122.6707, 21.vi.2013, coll. Darrell Ross, ex. Adelges tsugae on Tsuga heterophylla (1 larva [molecular voucher], YPM; ENT856663). Island County; Camano Island, 48.1474, –122.4411, 4.iv.2017, coll. Mark Whitmore, Isis Caetano, ex. Adelges tsugae on Tsuga heterophylla (1 larva, 2 puparia [molecular voucher], CUIC). King Co., Vashon, 47°27.200’N 122°26.950’W, 22.iii.2006,

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