Lasiopogon puyallupi (Cole & Wilcox, 1938) Mcknight & Cannings, 2020

Mcknight, Tristan A. & Cannings, Robert A., 2020, Molecular phylogeny of the genus Lasiopogon (Diptera: Asilidae) and a taxonomic revision of the bivittatus section, Zootaxa 4835 (1), pp. 1-115 : 71-74

Lasiopogon puyallupi stat. nov.

Lasiopogon willametti puyallupi Cole & Wilcox, 1938: 79 .

Diagnosis. ( Fig. 33 ) A medium-sized dark species from sandy rivers and beaches of northwestern Washington and British Columbia; all setae black except white setae of beard, coxae, femur, and tergite sides; thorax tomentum ranges from deep brown to grey with strong dorsocentral stripes; tergites usually with basal brown bands. Epandrial halves moderately concave medially, with black setae; phallus dorsal carina with protruding apex swollen but slightly dorsoventrally flattened, pointing in an acute angle with the aedeagal tube. Ovipositor dark brown/black, acanthophorite spines black.

Redescription. Body length ♂ 10.0– 11.1mm; ♀ 10.2–11.5mm. Head. HW ♂ 2.20–2.37mm; ♀ 2.16–2.48mm. FW ♂ 0.49–0.58mm; ♀ 0.52–0.57mm. VW ♂ 0.92–0.99mm; ♀ 0.93–1.05mm. VW/HW = ♂ 0.42–0.44; ♀ 0.41– 0.43. FW/VW = ♂ 0.51–0.59; ♀ 0.54–0.56. VD/VW = ♂ 0.12–0.16; ♀ 0.12–0.14. GH/GL = ♂ 0.40–0.45; ♀ 0.37–0.48. Face with silver-yellow tomentum; vertex with grey-brown tomentum, small bare patches between macroseta groups. Mystax, frontal, orbital, ocellar, and occipital setae black, occasionally a few ventral setae on mystax white; beard and labial setae white. Occipital macrosetae long and curved to right angle, longest (to 0.8mm) behind dorsolateral angle; lateral and ventral macrosetae shorter, straighter. Frontal and orbital setae fine and long; almost as long as ocellar macrosetae. Antennae. Black; setae black, no setae on postpedicel. Postpedicel long and rectangular; stylus broadly tapering. WPP/LPP = ♂ 0.26–0.31; ♀ 0.27–0.31. LAS/LPP ♂ 0.60–0.67; ♀ 0.54–0.59.

Thorax. Prothorax brown/grey, setae white; postpronotal lobes grey, lateral angle ferruginous, setae brown. Scutum tomentum ranges from dark brown to light grey. Dorsocentral stripes dark brown, cuticle subshining; acrostichal stripes slightly darker than base tomentum. All setae and macrosetae of scutum black; notal setae longer than scape. Dorsocentral macrosetae fine, mixed with other setae, 7–11 anteriors (to 1.0mm), 3–5 posteriors. Postalars 2–3, with a few shorter setae; supra-alars 2–4; presuturals 3; posthumerals 1–2. Scutellar tomentum grey-brown; rim inflated, leaving a semi-circular line impressed inside the dorsal edge; apical macrosetae black, very abundant (10+ on each side) and mixed with many other long setae. Pleural tomentum grey-tan. Katatergite macrosetae black, 8–10, with a few weaker setae; katepisternal setae moderately long, white; anepisternal setae black, 6–9 relatively strong (to 0.8mm) and mixed with other short setae along posterior margin, a patch of short setae on dorsal margin of sclerite; anepimeron with 0–2 short wispy setae. Legs. Base color black except apex of femur ferruginous; tomentum grey. No coxal peg. Macrosetae black, except basal 60% of femur; finer setae white on femur, black on tibia and tarsi. Ventral setae on femur as long as or slightly longer than femur width; dorsolateral macrosetae mixed with finer setae, 8–12 on profemur difficult to distinguish from surrounding finer setae, more prominent on mesofemur (4–8) and metafemur (11–17). Protibia with ventral macrosetae 2–3 times longer than tibial width; ventromedial patch of dense short setae on protibia pale yellow; dark brown on metatibia. Claws reddish brown over basal 60%, apically black. Wings. Veins dark brown; membrane faintly brown from microtrichia, darker around crossveins at apex of cells br and bm. DCI = 0.33–0.43. Halter cream, with no spot. Abdomen. Male. Cuticle dark brown/black; brown tomentum at bases of tergites, bands of grey tomentum cover apical 40% of each tergite as a straight band, extending anteriorly along lateral margins. Dorsal setulae dark brown; lateral setae all white; lateral macrosetae on first tergite mostly black (4–7), 1–3 straw-colored. Sternite tomentum grey; setae straw-colored/white. Female. As in male, but grey tomentum extends anteriorly along midline to divide brown into lateral spots, and lateral setae on tergites 4–7 black; sternite setae sparser than in male. Male genitalia. Epandrium and hypandrium/gonocoxite complex black; sparsely covered in thin grey tomentum and black setae, setal brush dark brown, dense. Epandrium elongate, the width of the halves in lateral view about 50% the length, widest in basal third; apex rounded with ventral tooth. In dorsal view, medial margins of epandrium moderately curved, curvature starts about 30% from base, outer margins gradually tapered; basal sclerite prominent. Phallus paramere sheath dorsally about 48% the length of phallus; paramere sheath without ventral ornamentation; dorsal carina protruding, with apex swollen but slightly dorsoventrally flattened and pointing in an imaginary acute angle with the aedeagal tube. Ejaculatory apodeme in lateral view relatively straight, with broad dorsal carina along entire length. Subepandrial sclerite with triangular central unsclerotized area in basal 35% narrowing to a parallel-sided gap in central portion; spines parallel-sided, blunt, densely distributed basally, apically, and along midline, sparse laterally.

Female genitalia. Undissected: Tergite 8 dark brown/black, with erect white setae; sternite 8 dark brown with short brown setae; lateral lobe setae strong, black; hypogynial valves with fine setae concentrated medially. Cerci brown/black with pale setae; acanthophorite spines black.

Type Material. LECTOTYPE (here designated) ♂ labelled: “[rectangular white label] Auburn, Wn./ V-10 1932 / J. Wilcox, Col.”; “[rectangular beige label] PARATYPE / Lasiopogon / willametti/ Cole & Wilcox”; “[rectangular beige label] FORM/ puyallupi/ Cole & Wilcox”; “[rectangular white label] JOSEPH WILCOX/ COLLECTION – 1981/ Gift to California / Academy of Sciences ”. Our lectotype label “[red and black bordered label] LECTOTYPE / Lasiopogon puyallupi / Cole & Wilcox 1938 / des. T.A. McKnight 2017” has been added. Deposited at the CAS.

PARALECTOTYPES (41 specimens designated): U.S.A.: WASHINGTON: King Co., Auburn [47.305 - 122.193], 10.v.1932, J. Wilcox (14♂ 1♀ CAS, 2♂ USNM, USNMENT1100230 – USNMENT1100231 ) ; Enumclaw [47.175 -122.029], 12.v.1932, J. Wilcox (10♂ CAS, 1♂ 1♀ [1 pr in cop] FISH) ; Pierce Co., Buckley [47.172 -122.039], 14.vi.1932, J. Wilcox (2♂ CAS) , 15.vi.1932, J. Wilcox (1♂ CAS) ; Mt. Rainier, Ipsut Creek Camp [46.9755 -121.828], 26.vi.1932, W.W. Baker (4♂ 1♀ CAS) ; Sumner [47.203 -122.248], 10.v.1932, J. Wilcox (1♂ CAS) , 17.vi.1932, J. Wilcox (1♂ CAS, 1♂ FISH, 1♀ USNM, USNMENT1100250 ) .

Other Material Examined (207 specimens). CANADA: BRITISH COLUMBIA: Agassiz [49.239 -121.767], 18.v.1927, H.H. Rose (1♀ CNC) ; Delta, Deas Island [49.125 -123.062], 26.iii.1983, R.A. Cannings (1♂ 1♀ RBCM, RBCM ENT 991-86355 , RBCM ENT 991-86358 ) , 6.iv.1984, R.A. Cannings (1♀ RBCM, RBCM ENT 991-86357 ) , 10.iv.1991, R.A. Cannings & H. Nadel (1♀ RBCM, RBCM ENT 991-67087 ) , 20.iv.1991, R.A. Cannings & H. Nadel (8♂ 1♀ RBCM, RBCM ENT 991-67067 , RBCM ENT 991-67070 , RBCM ENT 991-67079 – RBCM ENT 991-67081 , RBCM ENT 991-67083 , RBCM ENT 991-68073 – RBCM ENT 991-68074 , RBCM ENT 991-68084 ) , 12.v.1991, R.A. Cannings (1♂ 2♀ RBCM, RBCM ENT 991-67982 – RBCM ENT 991-67984 ) , 7.v.1997, R.A. Cannings (1♂ CUAC, 6♂ 1♀ RBCM, 1♂ 1♀ TAM) ; Dewdney, River Road South [49.144 -122.182], 19.v.1991, R.A. Cannings (1♂ RBCM, RBCM ENT 991-68073 ) ; Fraser River, mouth of Hunter Creek [49.359 -121.575], 4.v.1989, R.A. Cannings (1♀ RBCM, RBCM ENT 991-1377 ) ; Hope, Coquihalla River , 49.383 -121.45, 13.vi.1995, R.A. Cannings (1♂ RBCM) ; Hope, Hunter Creek, at Highway #1 bridge, under RR bridge [49.358 -121.574], 23.v.1996, S.G. Cannings (1♂ RBCM) ; Ladner [49.092 -123.095], 23.iv.1987, S.G. Cannings (2♂ CSUC, 1♂ RBCM), 18.v.1991, R.A. Cannings (7♂ 1♀ RBCM, RBCM ENT 991-68051 , RBCM ENT 991-68055 , RBCM ENT 991-68060 , RBCM ENT 991-68061 , RBCM ENT 991-68063 , RBCM ENT 991-68065 , RBCM ENT 991-68066 , RBCM ENT 991- 68068 ) , 19.v.1991, R.A. Cannings (1♂ RBCM, RBCM ENT 991-68054 ) ; Lillooet River, Meager Creek Hotsprings, terrace [50.576 -123.465], 19.vii.1988, C.S. Guppy (1♂ RBCM, RBCM ENT 991-8089 ) ; Lillooet River, near John Sandy Creek [50.486 -122.944], 9.vi.1981, R.J. Cannings (1♂ RBCM, RBCM ENT 991-86356 ) ; Meager Creek road, 16.v.1987, C.S. Guppy (1♂ RBCM) ; Miracle Beach Provincial Park [49.85 -125.094], 28.v.1988, S.G. Cannings (1♂ LACM, LACM ENT 334064 ) ; Tofino, Chesterman Beach [49.114 -125.893], 27.v.1994, R.A. Cannings (2♂ CSUC, 1♂ LACM, LACM ENT 334063 , 7♂ RBCM, RBCM ENT 010-002796 , 1♂ TAM) ; Tofino, Wickaninnish Beach [49.013 -125.675], 16.v.1994, R.A. Cannings (1♂ 2♀ RBCM) ; Vancouver Island, Brooks Peninsula, Cape Cook Lagoon , 50.2 -127.8, 2.viii.1981, R.A. & S.G. Cannings (1♂ 1♀ RBCM, RBCM ENT 991-86364 , RBCM ENT 991-86369 ) , R.A. Cannings (1♂ 6♀ [1 pr in cop] RBCM, RBCM ENT 991-86366 – RBCM ENT 991- 86371 ) ; Vancouver Island, Comox, Cape Lazo Road , 49.68828 -124.87444, 2.v.2006, R.A. Cannings & R.G. Bennett (1♂ 1♀ RBCM) .

U.S.A.: OREGON: Hood River Co., E Fork Hood River at Nottingham campground, 6 mi E of Mt. Hood peak, 45.36815 -121.57066, 15.vi.2013, T.A.McKnight (1♀ [1 EtOH] UMMZ) ; WASHINGTON: Clallam Co., Bogachiel River and Sol Duc River confluence, Hwy 110 jct Mora Rd , 47.91311 -124.539, 23.vi.2010, R.A. Cannings (1♂ 1♀ RBCM, RBCM ENT 011-012688 – RBCM ENT 011-012689 ) ; Bogachiel River, Highway 101 [47.892 -124.355], 31.v.1995, R.A. Cannings & H. Nadel (2♂ RBCM, 1♂ 1♀ TAM) ; Forks [47.96 -124.393], 2.vii.1920, E.P. VanDuzee (2♂ 1♀ EMEC) , 23.vii.1933, J. Wilcox (1♂ 2♀ CAS) ; Hoh River Campground, Hoh River , 47.85856 -123.93828, 24.vi.2010, R.A. Cannings (1♂ RBCM, RBCM ENT 011-012699 ) , 25.vi.2010, R.A. Cannings (1♀ RBCM, RBCM ENT 011-012714 ) ; Quillayute River County Park, trail to Quillayute River bank, 47.91194 -124.58283, 23.vi.2010, R.A. Cannings (1♂ RBCM, RBCM ENT 011-011681 ) ; Cowitz Co., N Fork Toutle River, 10 mi E of Castle Rock, 1 mi S Hwy 504 jct Tower Rd , 46.3477 -122.7053, 14.v.2015, T.A. McKnight (1♂ RBCM) ; Grays Harbor Co., W Fork Satsop River, Middle Satsop Road bridge, 47.04472 -123.52361, 30.v.1995, R.A. Cannings & H. Nadel (5♂ 1♀ RBCM) ; Jefferson Co., Hoh River Campground shore, upstream of Spruce Nature Trail , 47.86206 -123.92175, 25.vi.2010, R.A. Cannings (1♂ RBCM, RBCM ENT 011-011711 ) ; Hoh River Campground, Hoh River shoreline, 47.85856 -123.93828, 24.vi.2010, R.A. Cannings (6♂ 1♀ RBCM, RBCM ENT 011-012695 , RBCM ENT 011- 012697 – RBCM ENT 011-012698 , RBCM ENT 011-012700 – RBCM ENT 011-012702 ) , 25.vi.2010, R.A. Cannings (1♂ RBCM, RBCM ENT 011-011713 ) ; Hoh River jct Canyon Creek, Upper Hoh Road [47.81 -124.07], 1.v.1995, R.A. Cannings & H. Nadel (4♂ 1♀ RBCM) ; Hoh River rainforest, 7.vii.1968, W.W. Wirth (1♂ USNM, USNMENT1100761) ; Hoh River, Upper Hoh Road , 31.v.1995, R.A. Cannings & H. Nadel (9♂ 4♀ [1 pr in cop] RBCM) ; Hoh River, Upper Hoh Road 6.2 km E of Olympic NP boundary, 47.84272 -123.96406, 24.vi.2010, R.A. Cannings (1♂ RBCM, RBCM ENT 011-011691 ) ; Port Townsend [48.0825 -122.7622], 8.iv, A. Seaton (1♂ OSUC, OSUC 194508 ) ; Upper Hoh Road 7.9 km E of Olympic NP boundary, 47.85214 -123.95128, 22.vi.2010, R.A. Cannings (2♂ RBCM, RBCM ENT 011-011680 , RBCM ENT 011-012679 ) ; King Co., Auburn [47.305 -122.193], 10.v.1932, J. Wilcox (2♂ 6♀ [2 pr in cop] CAS, 2♂ 2♀ [1 pr in cop] EMEC) ; Enumclaw [47.175 -122.029], 12.v.1932, J. Wilcox (1♂ 1♀ [1 pr in cop] EMEC, 1♂ OSUC, OSUC 583677 ) ; Seattle [47.6064 -122.3308], 15.vii.1901, R.C. Osburn (1♀ OSUC, OSUC 194366 ) ; Lewis Co., Ashford 2.5 mi SE, Nisqually River jct Skate Cr Rd. , 46.7439 - 121.9796, 17.v.2015, T.A. McKnight (1♂ 1♀ [1 pr in cop] RBCM) ; Mason Co., Fernwood, Skokomish River beach at Skokomish River Road mile 4, 47.31792 -123.20542, 7.vi.2009, J. Miskelly (6♂ [3 EtOH] RBCM, RBCM ENT 010-002794 ) ; Pierce Co., Electron , 18.v.1936, J. Wilcox (1♀ CAS) ; Electron, Puyallup River , 47.0019 -122.1951, 6.vi.2009, J. Miskelly (1♂ RBCM) , 20.vi.2013, T.A. McKnight (1♂ [1 EtOH] UMMZ) , 11.vii.2014, T.A. McKnight (1♀ [1 EtOH] TAM) ; Mt. Rainier, Ipsut Creek Camp [46.9755 -121.828], 2.vii.1933, W.W. Baker (2♂ 1♀ CAS) , 23.vii.1935, S.E. Crumb (2♂ CAS) ; N. Puyallup [47.199 -122.263], 8.v.1932, C.H. Martin (1♂ 1♀ [1 pr in cop] FISH) ; Orting, Puyallup River , 47.08523 -122.20823, 4.vi.2009, J. Miskelly (2♂ RBCM, RBCM ENT 010-002792 , RBCM ENT 010-002795 ) ; Puyallup [47.199 -122.292], 2.v.1933, C.W. Getzendaner (1♂ CAS) , 11.v.1933, C.W. Getzendaner (1♀ CAS) , 13.v.1934, J. Wilcox (1♀ CAS) , 23.iv.1935, W.W. Baker (1♀ CAS) , 29.iv.1935, C.W. Getzendaner (1♀ CAS) ; Roy [47.005 -122.539], 29.iv.1932, R. Latta (2♂ CAS) ; Sumner [47.203 -122.248], 7.v.1932, C.H. Martin (1♂ ESUW) , 8.v.1932, C.H. Martin (1♂ 1♀ FISH, 1♀ ESUW), 10.v.1932, R. Latta (3♂ 2♀ CAS) , C.H. Martin (1♀ EMEC) , J. Wilcox (1♂ 1♀ [1 pr in cop] FISH, 1♀ OSUC, OSUC 583678 , 1♂ 1♀ RBCM, 1♀ USNM, USNMENT1100251 ), 11.v.1932, C.H. Martin (1♂ EMEC) , 12.v.1932, C.H. Martin (1♂ FISH, 1♂ LACM, LACM ENT 334065 ), 17.v.1932, C.H. Martin (1♂ 1♀ [1 pr in cop] FISH) , 20.vi.1932 (1♂ FISH) , 3.vi.1933, C.H. Martin (1♂ 1♀ FSCA) , 8.v.1936, W.W. Baker (2♂ CAS) , 16.v.1940, W.W. Baker (1♂ FSCA) ; Skagit Co., Edison [48.565 -122.443], 5.vi.1968, (1♀ WSUC) ; Thurston Co., Olympia, Pioneer Park, Deschutes River , 46.99304 -122.88711, 6.vi.2009, J Miskelly (2♀ RBCM, RBCM ENT 010-002793 ) ; Tumwater, Pioneer Park, Deschutes River , 46.994 -122.8881, 15.v.2015, T.A. McKnight (1♂ 1♀ [1 pr in cop] RBCM, 1♂ 1♀ [1 pr in cop] TAM, 3♂ [2 EtOH] UMMZ) .

Taxonomic Notes. Originally described as a variety of L. willametti in 1938. In accordance with ICZN Code article 45.6.4, this should be treated as a subspecific name, which we are now elevating to species. However, this subspecies was proposed merely for a subset of the paratypes of L. willametti and did not have an explicitly designated holotype. We have therefore interpreted the paratype specimens of L. willametti that also have labels for “Form puyallupi” as constituting a syntype series for the subspecies, and in the interest of stability, we are designating one of these specimens as a lectotype and all others with the “Form puyallupi” label as paralectotypes. Original paratypes of L. willametti without the “Form puyallupi” labels that nevertheless belong to L. puyallupi are listed in the Other Material Examined section, and not as paralectotypes.

Lasiopogon actius , L. puyallupi , and L. ripicola have mostly allopatric ranges but there is a curious tendency for polymorphic mystax color in the latter two species. We believe further work should investigate whether hybridization occurs in the peripheral contact zones for these three species.

Etymology. No explanation given in the original description, but evidently named for the type locality near Puyallup, Washington and the Puyallup River, despite Wilcox’s improper use of the singular masculine genitive (-i) instead of a locational adjectival suffix (-ensis).

Distribution ( Fig. 39 ) Nearctic; Canada: British Columbia and U.S.A.: Washington. Type locality: U.S.A., Washington: King Co., Auburn.

Phylogenetic Relationships. Member of the bivittatus section, actius group.

Natural History. Habitat: river banks and beach dunes ( Fig. 15 , likely also similar to Fig. 7 ). Perches on bare sand, cobble, fallen logs, or boulders; labels note “upper beach sand”, “ Carex / Elymus sand dunes”, “river edge sand and cobbles”, and “on log under bridge”. Dates collected: Mar 26 to Aug 2, most in May. One female with label indicating it was collected with prey pinned separately, but the prey item could not be found.

Figures

FIGURE 33. Matrix key of Lasiopogon morphology for species revised in this publication, indicating details of tergite tomentum, major setation, and dissected male genitalia. Taxa are grouped on pages with closely related and visually similar species. (33) L. actius, L. dimicki, L. puyallupi stat. nov.

FIGURES 37–40. Distribution maps of georeferenced records for Lasiopogon species revised in this publication, part 1 of 2. Each map corresponds to one page of the morphology tabular key.

FIGURES 14–21. Representative habitats of inland sand-perching Lasiopogon species collected for this revision. (14) Washington: Asotin Co., Snake River, sandy bank 4 mi S of Asotin jct Hwy 129, 21.iv.2014 (L. ripicola). (15) Washington: Thurston Co., Tumwater, Deschutes River @ Pioneer Park, 15.v.2015 (L. puyallupi stat. nov.). (16) Oregon: Linn Co., Crabtree Creek @ Hwy 226 bridge 8 mi N of Lebanon, 19.v.2012 (L. willametti). (17) Washington: Franklin Co., Juniper Dunes Wilderness, N entrance, 18 mi NE of Pasco, 20.iv.2014 (L. albidus). (18) California: Marin Co., San Antonio Creek, San Antonio Rd 4.5 mi S of Petaluma, 15.iii.2014 (L. zonatus, L. esau sp. nov.). (19) California: Riverside Co., Little San Gorgonio Creek @ Potrero blvd bridge, 2 mi NW of Beaumont, 9.iii.2014 (L. wilcoxi sp. nov.). (20) New Mexico: Grant Co., Gila River below Forks campsite, 6.v.2015 (L. karli sp. nov.). (21) California: Riverside Co., Santa Ana River at Mt. Rubidoux, 4.iii.2013 (L. drabicolum).

FIGURES 6–13. Representative habitats of coastal sand-perching Lasiopogon species collected for this revision. (6) Oregon: Lincoln Co., Gleneden Beach, NW tip of Salishan Spit, 17.v.2012 (L. actius on foredune, L. tumulicola sp. nov. in backdunes). (7) Washington: Grays Harbor Co., Westhaven State Park, dunes 2 mi NW of Westport, 20.vi.2013 (L. actius). (8) Oregon: Lincoln Co., Newport, Agate Beach State Park dunes, 21.vi.2013 (L. dimicki). (9) California: Humboldt Co., Clam Beach County Park ~5 mi SE of Trinidad, 15.v.2012 (L. canningsi sp. nov.). (10) California: San Mateo Co., Half Moon Bay State Beach, 5.vi.2013 (L. arenicola). (11) California: Monterey Co., Pacific Grove golf course, dunes in SW corner, 11.v.2012 (L. asilomar sp. nov.). (12) California: Santa Barbara Co., Rancho Guadalupe Dunes 1 mi S of parking lot, 9.v.2012 (L. littoris). (13) California: San Luis Obispo Co., Montaña de Oro State Park, dunes @ Sand Spit trailhead, 12.iii.2014 (L. bitumineus sp. nov., L. littoris weeks later).