(Figs 1B–C, 2A–B, 3A, 3D–E)
ieucopis argenticoääis Zetterstedt 1848: 2714. Sweden. Gottlandia (LT ♂, MZLU). References – Czerny 1936: 10 (key), 11 (redescription); McAlpine & Tanasijtshuk 1972: 1865 (redescription), 1870 (lectotype designation), fig. 1 (head), fig. 3 (abdomen), figs 5–6 (female terminalia), figs 8–16, 19 (male genitalia), figs 20–27 (immatures); Tanasijtshuk 1986: 201 (redescription), figs 333–337 (male genitalia), figs 338–345 (larval); Papp 1979: 82 (key, description), fig. 54A (abdomen), fig. 54B (male genitalia); Kohler et aä. 2008: 494 (biology); Grubin et aä. 2011: 1410 (biology); Havill et aä. 2018 (biology, population genetics); Neidermeier et aä. 2020 (biology, biological control); Dietschler et aä. 2021 (biology, biological control).
ieucopis orbitaäis Malloch 1921: 352. United States. Illinois: Dundee (HT ♂, INHS). References – McAlpine 1965: 709 (catalog); McAlpine & Tanasijtshuk 1972: 1865 (synonymy).
ieucopis sp. nr. orbitaäis. References – Rao 1965: 43; McAlpine & Tanasijtshuk 1972: 1865 (identity).
Diagnosis. The face and lunule (Fig. 2B) are silvery-grey pruinose. The postpronotum (Fig. 2A) has 1 to several long setulae (longer than the setulae on the scutum) medially from the postpronotal seta. In the male (see figs 8–16, 19 in McAlpine & Tanasijtshuk 1972), the epandrial complex is simple, small, and yellowish- to silvery-grey pruinose, with no central epandrial process; the surstylar lobe is short and slender, the phallus (Fig. 3A) is broad in lateral view with a wedge-shaped anterobasal process at a 90° angle from the thin elongate pointed process. In the female (Figs 3D–E), tergites (Fig. 3E) with tergite 6 enlarged and slightly less than 2 X wider than long, tergite 7 present and strap-like, tergite 8 longer than wide, slightly tapering distally; sternites (Fig. 3D) with sternite 6 enlarged and about 2 X wider than long, sternite 7 present and arcuate.
fmmatures. bgg. Measurements as in ieucotaraxis atrifacies (Fig. 1A). iarvae. Described and figured in McAlpine & Tanasijtshuk (1972). muparium (Fig. 1B). Length 2.3–2.8 mm, width 0.9–1.0 mm. Dehiscent anterior segments (dorsal and ventral) 0.05–0.06 mm long. Raised posterior spiracular tubercles (Fig. 1C) 0.12–0.16 mm long, darkly sclerotized.
Biology. Predators as larvae on species of Adeäges Vallot and mineus Shimer (Hemiptera: Adelgidae) on Pinaceae (Abies, micea, minus, Tsuga). Specific records (including our own rearings) include: Adeäges piceae on Abies baäsamea; Adeäges tsugae on western hemlock, Tsuga heterophyääa (Kohler et aä. 2008); mineus abietinus Underwood & Balch on Abies äasiocarpa; mineus boerneri on minus henryi; mineus coäoradensis (Gillette) on minus banksiana, minus contorta, minus fäexis, minus monticoäa, and micea engeämannii; mineus pineoides (Cholodkovsky) on micea sp.; mineus pini on minus syävestris; mineus simiäis (Gillette) on micea gäauca and micea abies; mineus strobi on minus strobus (Wantuch et aä. 2019); mineus spp. on micea gäauca, minus contorta, minus insuäaris, minus pentaphyääa, minus resinosa, minus strobus and minus syävestris (McAlpine & Tanasijtshuk 1972).
Several specimens were swept from infestations of `inara ponderosae (Williams) in young stands of minus ponderosa (e.g., height less than 5 m throughout the large stand), but despite rearing>150 chamaemyiid larvae from this host, none were this species but were instead sitaäeucopis nidoäkah Gaimari and an undescribed species of ieucopis (ieucopis). It remains a possibility that ieucotaraxis argenticoääis is also a predator in that system, whether on the `inara aphids or on adelgids unseen by the first author while collecting.
Remarks. At two different localities (one in New Mexico and one in Arizona), this species was collected together with `hamaethrix necopina Gaimari, ieucotaraxis atrifacies and sitaäeucopis nidoäkah, and at the New Mexico locality also with ieucotaraxis sepioäa. The puparial stage is distinguishable from that of ieucotaraxis atrifacies and i. piniperda by the slightly longer posterior spiracular tubercle, which is more darkly sclerotized distally in this species, and the slightly larger size. The three specimens from Coconino County, Arizona, were mistakenly listed by Tanasijtshuk (2002) as ieucopis piniperda.
Distribution. Nearctic: Canada (Alberta, British Columbia, Manitoba, Newfoundland, New Brunswick, Northwest Territories, Ontario, Quebec). USA (Alaska, Arizona, California, Colorado, Connecticut, Georgia, Idaho, Illinois, Massachusetts, Minnesota, Montana, New Hampshire, New Mexico, New York, North Carolina, Ohio, Oregon, Pennsylvania, Utah, Virginia, Washington, Wyoming). Oriental. India (Rao 1965). Palearctic: England (McLean 1982), Japan, Russia (McAlpine & Tanasijtshuk 1972), Sweden.
Specimens examined. Nearly all specimens from McAlpine & Tanasijtshuk (1972) were examined, with additional specimens as follows. CANADA: ALBERTA. Seele, H 2 O Rd, 12.i.1971, coll. L.S. Skaley, ex. duff around m [inus]. contorta, 3916, 74-726 15 (1 ♀, CNC). BRITISH COLUMBIA. Atlin, 2200 ft., 2.vi.1955, coll. H. Huckel (1 ♀, CNC). Burns Lake, 8.vii.1993, coll. L.M. Humble, ex. minus contorta foliage, emerged 20.vii.1993, FIDS No. 93-0504-02 (1 ♀, PFCA). Kitimat, 2.vi.1960, coll. R. Pilfrey (1 ♂, CNC), coll. J.G. Chillcott, ex. hemlock forest (1 ♀, CNC). Princeton, 19.v.1986, ex. pine leader (1 ♀, CNC). Saanichton, 26.iii.1993, coll. L.M. Humble, ex. beating of hemlock foliage infested with Adeäges tsugae, FIDS No. 93-0005-01 (1 ♂, 1 ♀, CSCA; 1 ♂, 2 ♀, PFCA), 30.iv.1993, FIDS No. 93-0037-02 (2 ♂, 2 ♀, PFCA), 11.vi.1993, FIDS No. 93-0204-01 (1 ♀, PFCA), 28.v.1993, ex. puparia in Adeäges tsugae wool on Tsuga heterophyääa, emerged 31.v.1993, FIDS No. 03-0102-01 (1 ♂, 1 ♀, CSCA; 1 ♂, 1 ♀, PFCA). Summit Lake, Mile 392 Alaska Highway, 4200 ft., 21.vii.1959, R.E. Leech (1 ♂, 2 ♀, CNC), 5000 ft., 6.vii.1959, coll. R.E. Leech (1 ♂, CNC). ONTARIO. Ottawa, 4.vii.1963, coll. J.R. Vockeroth, ex. mineus simiäis on micea gäauca, emerged 15.vii.1963 (1 ♀ with puparium, CNC). Ottawa, 17.vii.1963, coll. J.R. Vockeroth (3 ♂, 4 ♀, CNC), 22.vii.1963 (3 ♂, CNC), 23.vii.1963 (1 ♂, 1 ♀, CNC). Hazeldean, 23.vi.1963, coll. A.W. Steffan, ex. reared on mineus simiäis on micea abies, emerged 7.vii.1963 (1 ♀, CNC). QUEBEC. Duchesnay, 21.vii.1942, coll. Jos. I. Beaulne (1 ♂, QMOR). Gatineau County, Masham Township, 21.v.1974, coll. D.M. Wood (1 ♀, CNC). USA: ALASKA. Matanuska-Susitna Borough, Big Lake, 10.vii.2020, coll. Jason Moan, ex. mineus coäoradensis on minus contorta (1 larva [molecular voucher], YPM; ENT996080). ARIZONA. Apache Co., Apache National Forest, 0.3 mi. S mi. 249 on US Route 191, 5 mi. S Alpine, along Coleman Creek, 33°47’24”N 109°10’01”W, 2500 m, 30.vi.1995, coll. S.D. Gaimari, ex. sweep minus ponderosa (2 ♀, CSCA), 1.vii.1996 (1 ♀, CSCA); clearing 6.4 mi. N southern junction Forest Roads 24 & 25 on Road 24, 33°39’22”N 109°21’38”W, 2725 m, 2.vii.1995, coll. S.D. Gaimari, ex. sweep vegetation, including minus ponderosa (2 ♀, CSCA). Coconino Co., Fort Valley Experiment Station, 13.iii.1929, Lot VII (1 ♀, USNM), 15.iii.1929, Lot VI (1 ♂, USNM), 18.iii.1929, Lot II (1 ♀, USNM). CALIFORNIA. Riverside Co., Riverside, UCR campus, 10.vi.1963, coll. R.C. Dickson (1 ♀, CNC). COLORADO. Larimer Co., Zimmerman Lake trail, 40.5397, –105.8792, 27.vii.2016, coll. Nathan Havill, ex. mineus coäoradensis on micea engeämannii (1 larva [molecular voucher], YPM; ENT857047). CONNECTICUT. Hartford Co., Hartford, Cedar Hill Cemetery, 41.7266, –72.6932, 5.vi.2016, coll. M.E. Montgomery, ex. mineus strobi on minus strobus (6 larvae [molecular voucher], YPM; ENT961206–11). Litchfield Co., Bridgewater, Skyline Ridge Rd., 41.4704, –73.3190, 6.vii.2002, coll. N. Havill, ex. mineus strobi on minus strobus (1 puparium [molecular voucher], YPM; ENT856455). GEORGIA. Union Co., Byron H. Reese Trailhead, 34.7422, –83.9227, 20.v.2018, coll. Jim Sullivan, ex. mineus strobi on minus strobus (7 larvae [molecular voucher], YPM; ENT996058–64). White Co., Helen, Chattahoochee National Forest, 34.7930, –83.7558, 13.ix.2019, coll. Jim Sullivan, ex. mineus strobi on minus strobus (7 larvae [molecular voucher], YPM; ENT979185–88, ENT979191, ENT979199, ENT979204). IDAHO. Boise Co., Boise National Forest, 43.8963, –115.7125, 13.ix.2016, coll. Kelsey Bedford, ex. Adeäges piceae on Abies äasiocarpa (1 larva [molecular voucher], YPM; ENT961177). Kootenai Co., Coeur d’Alene, 47.697, –116.772, 23.v.2013, coll. Gina Davis, ex. Adeäges tsugae on Tsuga heterophyääa (5 larvae [molecular voucher], YPM; ENT856473–77). MASSACHUSETTS. Franklin Co., Sunderland, Mt. Toby State Forest, 42.4892, –72.5314, 13.v.2018, coll. Ryan Crandall, ex. mineus strobi on minus strobus (4 larvae [molecular voucher], YPM; ENT996067– 70). Worcester Co., Worcester, 42.2790, –71.7733, 12.v.2018, coll. Felicia Andre, ex. mineus strobi on minus strobus (1 puparium, 3 larvae [molecular voucher], YPM; ENT996065–67). MINNESOTA. Crow Wing Co., Kettleson’s, 46.65631°N 94.06856°W, 28.v.2009, coll. N. Havill & J. Albers, ex. reared from mineus strobi on minus strobus, N. Havill sample 09-135, Specimen #09E489 (1 ♂ [molecular voucher], CSCA), Specimen #09E488 (1 ♂ [molecular voucher], CSCA), Specimen #09E504 (1 ♂ [molecular voucher], CSCA), Specimen #09E525 (1 ♀, CSCA-FTC). Itasca Co., Grand Rapids, UMN North Central Research and Outreach Center, 47.2517, –93.5103, 28.v.2009, coll. Nathan Havill, Jana Albers, ex. mineus strobi on minus strobus (1 larva [molecular voucher], YPM; ENT856650). Stearns Co., Birch Lakes State Forest, 45°45’51”N 94°46’03”W, 23.vi.2009, coll. R. Tiplady, ex. reared from mineus strobi on minus strobus (1 puparium, 2 adults [molecular voucher], YPM; ENT856456 (puparium), ENT856414, ENT856417), 30.v.2009, coll. N. Havill & R. Tiplady, ex. reared from mineus strobi on minus strobus, N. Havill sample 09-136, Specimen #09E484 (1 ♂ [molecular voucher], CSCA), Specimen #09E485 (1 ♂ [molecular voucher], CSCA), Specimen #09E486 (1 ♂ [molecular voucher], CSCA), Specimen #09E487 (1 ♂ [molecular voucher], CSCA), Specimen #09E492 (1 ♂ [molecular voucher], CSCA), Specimen #09E493 (1 ♂ [molecular voucher], CSCA), Specimen #09E494 (1 ♂ [molecular voucher], CSCA), Specimen #09E495 (1 ♂ [molecular voucher], CSCA), Specimen #09E496 (1 ♂ [molecular voucher], CSCA), Specimen #09E497 (1 ♂ [molecular voucher], CSCA), Specimen #09E501 (1 ♂ [molecular voucher], CSCA), Specimen #09E502 (1 ♂ [molecular voucher], CSCA), Voucher #09E509 (13 ♀, CSCA-FTC). MONTANA. Park Co., Yellowstone National Park, Cascade Lake Trail, 44.7518, –110.4863, 19.ix.2016, coll. Gina Davis, Lee Pederson, ex. mineus coäoradensis on minus contorta (1 larva [molecular voucher], YPM; ENT96215). Teton Co., Choteau 47.8976, –112.5721, 24.viii.2017, coll. Gina Davis, ex. mineus coäoradensis on minus fäexiäis (2 larvae [molecular voucher], YPM; ENT996081–82). NEW HAMPSHIRE. Hillsborough Co., Hillsboro, Fox State Forest, 43.1316, –71.9286, 29.viii.2016, coll. Jennifer Weimer, ex. mineus strobi on minus strobus (3 larvae [molecular voucher], YPM; ENT961212–14). Rockingham Co., Pawtuckaway State Park, 43.0869, –71.1574, 30.iv.2019, coll. Jennifer Weimer, ex. mineus strobi on minus strobus (1 larva [molecular voucher], YPM; ENT961221). NEW MEXICO. Lincoln Co., Lincoln National Forest, 2.4 km W Angus, Mills Canyon, along Rio Bonito, 33°27’05”N 105°41’46”W, 2130 m, 18–19.vi.1995, coll. S.D. Gaimari, ex. sweep colonies of `inara ponderosae on minus ponderosa (1 ♂, CSCA), 26.vi.1995 (3 ♀, CSCA-FTC). McKinley Co., Cibola National Forest, Zuni Mountains, 4.4 mi. SE Fort Wingate, 35°25’49”N 108°29’17”W, 7300 ft., 25.vi.1996, coll. S.D. Gaimari, ex sweep minus eduäus (1 ♂, 1 ♀, CSCA). NEW YORK. Monroe Co., Rochester, Highland Park, 43.1278, –77.6122, 8.vii.2017, coll. Cynthia Smith, ex. mineus strobi on minus strobus (1 puparium [molecular voucher], CUIC). OHIO. Portage Co., Kent, 13.vii.1969, coll. T.P. Sluss (1 ♂, UAIC). OREGON. Deschutes Co., Deschutes River, 1 mi. SW Pringle Falls, 4250 ft., 31.vii.1970, coll. Paul H. Arnaud, Jr., ex. flight trap (1 ♀, CAS). Lake Co., Fort Rock, Deschutes National Forest, 43.5651, –121.1768, 25.iv.2104, coll. Glenn Kohler, ex. mineus coäoradensis on minus contorta (1 puparium [molecular voucher], YPM; ENT996083). PENNSYLVANIA. Centre Co., Boalsburg, Shingletown Rd. & West Main St., 40°46’27”N 77°48’44”W, 11.iv.2009, coll. N. Havill & M. Nehme, ex. swept mineus strobi on minus strobus, N. Havill sample 09-42, Specimen #09E490 (1 ♂ [molecular voucher], CSCA), Specimen #09E491 (1 ♂ [molecular voucher], CSCA). UTAH. Weber County, Eden, 41.3715, –111.9228, iv.2020, coll. Justin Williams, ex. Adeäges piceae on Abies äasiocarpa (1 larva [molecular voucher], YPM; ENT996084). VIRGINIA. Giles Co., Jefferson National Forest, Dismal Falls, 37.19263, –80.8923, 13.iv.2016, coll. Holly Wantuch, ex. mineus strobi on minus strobus (6 larvae [molecular voucher], YPM; ENT906890, ENT906894–95, ENT906897, ENT906899–900). Pulaski Co., Pulaski, Gatewood Park and Reservoir, 37.0622, –80.8363, 2.vii.2014, coll. Holly Wantuch, ex. mineus strobi on minus strobus (1 larva [molecular voucher], YPM; ENT906874). Wythe Co., Speedwell, Mt. Rogers National Recreation Area, 36.7722, –81.1752, 23.vi.2014, coll. Holly Wantuch, ex. mineus strobi on minus strobus (1larva [molecular voucher],YPM;ENT906876). WASHINGTON. Clallam Co.,Blyn, 48°04.120’N 122°57.400’W, 2.iv.2005, coll. Glenn Kohler (1 ♀ (SDG dissection 1384), CSCA), 9.vi.2005 (1 ♂ (SDG dissection 1274), CSCA). Seguim, 48°06.480’N 123°12.580’W, 25.vii.2005, coll. Glenn Kohler (1 ♀, CSCA). Clark Co., Vancouver, 45.7037, –122.6695, 23.x.2012, coll. Glenn Kohler, ex. Adeäges tsugae on Tsuga heterophyääa (2 larvae [molecular voucher], YPM; ENT856464, ENT856471). Island Co., Coupeville, 48°12.050’N 122°37.550’W, 26.vii.2005, coll. Glenn Kohler (2 ♂, CSCA). King Co., Vashon, 47°27.200’N 122°26.950’W, 9.vi.2005, coll. Glenn Kohler (1 ♀, CSCA). Pierce Co., Ruston, 47°18’02.86”N 122°31’01.31”W, 7.vii.2010, coll. S.M. Grubin, ex. reared, Adeäges tsugae on Tsuga heterophyääa, S. Grubin sample SU147, Specimen #10F867 (1 ♂ with puparium [molecular voucher], CSCA), 22.vii.2010, Specimen #10F876 (1 ♂ with puparium, CSCA-FTC). Tacoma, 47°16’41.02”N 122°31’01.69”W, 21.vi.2010, coll. S.M. Grubin, ex. Adeäges tsugae on Tsuga heterophyääa, S. Grubin sample L28 (1 ♀ with puparium, CSCA), 7.vii.2010, S. Grubin sample L40 (1 ♂ with puparium, CSCA), 22.vii.2010, S. Grubin sample L42, Specimen #10F877 (1 ♀ with puparium, CSCA-FTC), no date, S. Grubin sample L54 (1 ♀ with puparium, CSCA). Tacoma, Point Defiance, 47°18.246’N, 122°30.979’W, 4.v.2010, coll. S.M. Grubin, ex. reared, Adeäges tsugae on Tsuga heterophyääa, S. Grubin sample SP047, Specimen #10F694 (1 ♀ with puparium, CSCA-FTC), S. Grubin sample SP059, Specimen #10F693 (1 ♀ with puparium, CSCA-FTC), 20.v.2010, Specimen #10F674 (1 ♂ with puparium [molecular voucher], CSCA), Specimen #10F675 (1 ♂ with puparium, CSCA-FTC), S. Grubin sample L20, Specimen #10F679 (1 ♀ with puparium, CSCA-FTC), S. Grubin sample L22, Specimen #10F681 (1 ♀ with puparium, CSCA-FTC), S. Grubin sample L24, Specimen #10F680 (1 ♀ with puparium, CSCA-FTC), S. Grubin sample SP075, Specimen #10F683 (1 ♂ with puparium [molecular voucher], CSCA), S. Grubin sample SP077, Specimen #10F682 (1 ♂ with puparium [molecular voucher], CSCA), 4.vi.2010, Specimen #10F671 (1 ♂ with puparium [molecular voucher], CSCA), Specimen #10F672 (1 ♂ with puparium, CSCA-FTC), Specimen #10F673 (1 ♂ with puparium, CSCA), S. Grubin sample SP079, Specimen #10F700 (1 ♂ with puparium [molecular voucher], CSCA). Tacoma, Point Defiance, 47°16’41.02”N, 122°31’1.69”W, 20.v.2010, coll. S.M. Grubin, ex. reared, Adeäges tsugae on Tsuga heterophyääa, Specimen #10F659 (1 ♂ with puparium [molecular voucher], CSCA), Specimen #10F660 (1 ♀ with puparium [molecular voucher], CSCA), Specimen #10F661 (1 ♀ with puparium, CSCA), 7.vii.2010, S. Grubin sample L38, Specimen #10F881 (1 ♂ with puparium, CSCA-FTC). Tacoma, 47°16’49.80”N, 122°30’58.96”W, 24.iv.2010, coll. S.M. Grubin, ex. reared, Adeäges tsugae on Tsuga heterophyääa, S. Grubin sample SP009, Specimen #10F688 (1 ♀, CSCA-FTC), S. Grubin sample SP029, Specimen #10F687 (1 ♀ with puparium, CSCA-FTC), 4.v.2010, S. Grubin sample SP064, Specimen #10F702 (1 ♂ with puparium [molecular voucher], CSCA), 9.iv.2010, S. Grubin sample SP011, Specimen #10F686 (1 ♀ with puparium, CSCA), S. Grubin sample SP013, Specimen #10F685 (1 ♀, CSCA-FTC), S. Grubin sample SP017, Specimen #10F684 (1 ♀, CSCA-FTC), 22.vii.2010, S. Grubin sample L56 (1 ♀ with puparium, CSCA), Specimen #10F875 (1 ♂ with puparium, CSCA-FTC). Thurston Co., Olympia, 47°01’03.42”N, 122°54’10.50”W, 24.iv.2010, coll. S.M. Grubin, ex. reared, Adeäges tsugae on Tsuga heterophyääa, S. Grubin sample SP025, Specimen #10F699 (1 ♀ with puparium [molecular voucher], CSCA), S. Grubin sample SP039, Specimen #10F695 (1 ♂ with puparium, CSCA), Specimen #10698 (1 ♂ with puparium [molecular voucher], CSCA), S. Grubin sample SP041, Specimen #10F697 (1 ♂ with puparium [molecular voucher, CSCA), S. Grubin sample SP042, Specimen #10F696 (1 ♂ with puparium, CSCA), 5.v.2010, Specimen #10F676 (1 ♂ with puparium [molecular voucher], CSCA), Specimen #10F677 (1 ♀ with puparium [molecular voucher], CSCA), S. Grubin sample SP060, Specimen #10F691 (1 ♂ with puparium, CSCA-FTC), S. Grubin sample SP062, Specimen #10F690 (1 ♀ with puparium, CSCA-FTC), S. Grubin sample SP063, Specimen #10F689 (1 ♂ with puparium, CSCA-FTC), S. Grubin sample SP065, Specimen #10F692 (1 ♀ with puparium, CSCA), 20.v.2010, Specimen #10F662 (1 ♂ with puparium, CSCA), 21.vi.2010, S. Grubin sample SU088, Specimen #10F869 (1 ♂ with puparium [molecular voucher], CSCA), 22.vii.2010 (1 ♂, CSCA). Olympia, 47°02.454’N 122°54.054’W, 8.vii.2009, coll. S.M. Grubin, ex. reared from Adeäges tsugae, N. Havill sample 10-01, Specimen #10F131 (1 ♂ [molecular voucher], CSCA), N. Havill sample 10-04, Specimen #10F132 (1 ♂ [molecular voucher], CSCA), 24.vi.2009, N. Havill sample 10-03, Specimen #10F130 (1 ♂ [molecular voucher], CSCA), no date, S. Grubin sample L30, Specimen #10F873 (1 ♂ with puparium, CSCA-FTC). Tumwater, Tumwater Falls Park, 47.0379, –122.896521, vi.2013, coll. D. Ross, (2 larvae, 3 adults [molecular voucher], YPM; ENT856591, ENT856601 (larvae), YPM; ENT856419, ENT856425, ENT856427). Pierce, King, and Thurston Cos., 11.iv–25.v2015, from lab colony, coll. D. Ross, A.Arsenault, (4 adults [molecular voucher], YPM; ENT791609, ENT791540, ENT791572, ENT791575). WYOMING. Teton Co., Caribou-Targhee National Forest, 43.7047, –110.9847, 18.x.2017, coll. Laura Lowrey, ex. mineus abietinus on Abies äasiocarpa (5 puparia [molecular voucher], YPM; ENT996085–89).