Iratiquedius prostans (Horn, 1878) Brunke, 2022

Brunke, Adam J., 2022, A world generic revision of Quediini (Coleoptera, Staphylinidae, Staphylininae), part 1. Early diverging Nearctic lineages, ZooKeys 1134, pp. 129-170 : 129

Iratiquedius prostans (Horn, 1878) comb. nov.

Figs 2E , 3F , 6L-P , 9C , 11B (map)

Quedius prostans Horn, 1878: 165.

Quedius rupimontis Casey, 1915: 418.

Quedius (Raphirus) prostans : Smetana 1971 (redescription); Brunke et al. 2016, 2019, 2021 (phylogeny, outside of Raphirus ).

Type locality.

‘California’, United States.

Type material.

Lectotype (male, MCZ): The lectotype of this common, widespread species was not examined as its identity was not in doubt.

Non-type material.

Canada: British Columbia: Central Kootenay: 8 mi W Creston, 10.VI.1968, Campbell and Smetana (8, CNC); Columbia-Shuswap: Mount Revelstoke National Park, 600 m, 17.VIII.1971, J.M. Campbell (1, CNC); Fraser Valley: 7 mi W Hope, 3.VI.1968, Campbell and Smetana (1, CNC); Squamish-Lillooet: Garibaldi Provincial Park, Diamond Head Trail, 1128 m, 1.VIII.1975, J.M. Campbell & B.A. Campbell (3, CNC); Garibaldi Provincial Park, Mimulus Creek, 1645 m, 8.VIII.1975, J.M. & B.A. Campbell (1, CNC); Mount Garibaldi, 30.V.1968, Campbell and Smetana (22, CNC); Vancouver Island: Elk Lake trail, 48.534470, -123.398647, 27.IX.2020, A. Davies, sedge litter at base of cottonwood, edge of lake (1, CNC); Goldstream Park, 27.Vl1968, A. Smetana (1, CNC); Goldstream Park, 5 mi N Victoria, 27.V.1968, Campbell and Smetana (21, CNC); same except, 6.VI.1975, JM & BA Campbell (3, CNC); Gabriola, sifting moss along pond edge, 3.V.1994, Lot 3, BF & JL Carr (1, CNC); same except Lot4, under wood on wet muck near pond (1, CNC); Nitinat Lake, at Caycuse River, 21.VI.1989, sifting wash up on beach, Lot 5, BF & JL Carr (2, CNC); Duncan, Mount Tzouhalem, 19.X.2008, A. Davies (2, CNC); Lake Cowichan, South Shore Road, 2.3 km N of town, wet moss, 16.VI.1979, I. Smith (9, CNC); Lake Cowichan, spring run beside North Shore Road, 1.7 km N town, moss and litter, 7.VI.1979, I. Smith (5, CNC); 10 mi E [Port] Alberni, MacMillan Provincial Park, 26.V.1968, Campbell and Smetana (1, CNC); near Mount Finlayson Trail, Malahat, Goldstream Provincial Park, moss on rock, 11.VII.1979, I. M. Smith (4, CNC); Port Alberni, Mount Arrowsmith, nr. road to ski area, 11.6 km off Highway 4, 28.VI.1979, I.M. Smith, moss on rocks and sticks in stream (5, CNC); Hillcrest Rd., 16 km S Mesachie Lake, along Lens Creek, 12.VII.2010, A. Davies (1, CNC).

United States: Arizona: Apache Co.: Chuska Mountains, Wagonwheel Campground, sifting leaf litter, 2250 m, 12.VII.1976, J.M. Campbell (5, CNC); same except: sifting moss along waterfall (28, CNC); California: Butte Co., 3 mi NE Loma, 3.V.1981, sift litter along spring, DS Chandler (1, FMNH); Mountain House [Brush Creek], 7.V.1981, sift litter along spring, DS Chandler (1, FMNH); Calaveras Co.: Big Trees State Park, 38.2775, -120.310556, 25.V-26.VI.2010, FIT, A.R. Cline & S.L. Winterton (12, UTCI); El Dorado Co.: 5 mi SW Kyburz, 1219 m, 6.V.1968, Campbell & Smetana (23, CNC); same except: 7 mi E Kyburz (6, CNC); Glenn Co.: 5 mi NE Alder Spring, 20.IX.1979, sift oak litter along spring, DS Chandler (2, FMNH); Humbolt Co.: Garberville, Garberville-Harris Road, 5-6 miles east of Garberville, 13.VII.1965, under stones and pieces of sod, Lot 2, BF & JL Carr (1, CNC); Los Angeles Co.: Mount Wilson, 600 m, 26.V.1978, J.O. Martin (1, CNC); Marin Co.: Point Reyes National Seashore, 2 mi W Inverness, 22.III.1983, A. Smetana (3, CNC); Mendocino Co.: Mendocino, 24.VI.1954, Helfer (2, CNC); Placer Co.: 4 mi S Truckee, Truckee River, 5.V.1968, Campbell and Smetana (1, CNC); Lake Tahoe, Tahoe City, 1950 m, 7.VII.1986, A. Smetana (1, CNC); San Bernardino Co.: San Bernardino Mountains, 1 mi NE Angelus Oaks, Cold Creek at Highway 38, 1828 m, 12.III.1983, A. Smetana (33, CNC); San Bernardino Mountains, Highway 38, 3 mi SW Onyx Summit, 2346 m, 14.III.1983, A. Smetana (13, CNC); San Diego Co.: Laguna Mountains, Little Laguna Lake, 5.III.1983, A. Smetana (10, CNC); Mount Laguna, Carex clumps at stream, 25.IX.1981, JM Campbell (3, CNC); Mount Palomar, 1524 m, sifting leaf litter, 27.IX.1981, J.M. Campbell (2, CNC); Mount Palomar, Fry Trail Campground [Fry Creek], 8.III.1983, A. Smetana (1, CNC); San Francisco Co.: San Francisco, 30.V.1911, Van Dyke (4, CNC); Siskiyou Co.: Calahan, Lillypad Lake, old rotten logs, 8.VII.1991, Lot 1,BF & JL Carr (1, CNC); McBride Springs, Mount Shasta, 1447 m, 20.VI.1974, A&D Smetana (1, CNC); Trinity Co.: 10 mi N Junction City, 762 m, 10.VII.1979, J.M. & B.A. Campbell (4, CNC); 12 mi N Junction City, 1030 m, 13.VII.1979, J.M. & B.A. Campbell (4, CNC); Upper Canyon Creek Meadows, 1463 m, 13.VII.1979, J.M. & B.A. Campbell (2, CNC); 19 mi W Coffee Creek Station, Shasta National Forest, 1219 m, 14.VII.1979, J.M. & B.A. Campbell (1, CNC); 4 mi W Forest Glen, 9.VII.1979, J.M. & B.A. Campbell (2, CNC); Tulare Co.: Wishon Campground, 12 mi NE Springville, Meadow Creek, sifting washed up debris in fast flowing creek, 21.VI.1993, Lot 11 BF & JL Carr (4, CNC); 28 mi NNW Kernville, Thompson Camp Spring, 1676 m, 30.V.1981, L. Herman (1, CNC); Tuolumne Co.: Strawberry, 3.VIII.1960, DG Cavanaro (3, CNC); Idaho: Boise Co.: 10 mi NE Idaho City, 10 Mile Campground, sifting moss, 18.VII.1981, J.M. Campbell (23, CNC); Elmore Co.: Boise National Forest, Ice Springs, 1463 m, 21.VII.1981, J.M. Campbell (1, CNC); middle fork of Boise River and Dutch Creek, 1370 m, sifting moss, 19.VII.1981, J.M. Campbell (20, CNC); Nevada: Douglas Co.: Zephyr Cove, 1900 m, 9.VII.1986, A. Smetana (13, CNC); New Mexico: Sandoval Co.: Sandia Mountain, Cibola National Forest, Las Huertas Creek, wet-soaked moss encrusted with calcareous deposits, springy slope, 8.VII.1981, A. Smetana (11, CNC); Oregon: Benton Co.: Mary’s Peak, 1158 m, 27.VII.1979, J.M. & B.A. Campbell (1, CNC); Mary’s Peak, waterfalls, 1066 m, 5.V.1973, E.M. Benedict (1, CNC); Clackamas Co.: Camp Creek, 3.5 mi SE Rhododendron, 700-730 m, 27.VI.1974, A&D. Smetana (2, CNC); Timberline Lodge Road, Mt. Hood, 28.VI.1974, A&D Smetana (1, CNC); Coos Co.: Dune Park, 3 mi N and 2 mi W North Bend, sunny frost pockets, 15.I.1972, E. M. Benedict (1, CNC); Curry Co.: Agness Rd., [crossing at Wake Up Rilea Creek, under stones and in little pools of water along shady, cascading creek, 10.VIII.1978, B.F. & J.L. Carr] (4, CNC); Deschutes Co.: 12 mi SW Sisters, 1341 m, J.M. & B.A. Campbell, 23.VII.1979 (11, CNC); Douglas Co.: 27.3 miles NE Reedsport, at Smith River Falls, 29.VI.1978, L & N Herman (2, CNC); Grant Co.: Dixie Summit, Highway 26, 1615 m, sifting moss, 22.VII.1981, J.M. Campbell (71, CNC); Malheur National Forest, 2 km NW Highway 26, Forest Road 1218, 1670 m, sifting moss, 22.VII.1981, JM & BA Campbell (13, CNC); Malheur National Forest, 7 km NW Highway 26, 2040 m, sifting old pile of hay, 22.VII.1981, JM & BA Campbell (2, CNC); road 2610, below Dixie Butte, 2050 m, 2.VI.1989, A. Smetana (8, CNC); Strawberry Range, Strawberry Campground, 1780 m, 1.VI.1989, A. Smetana (1, CNC); Strawberry Range, road 650, Fawn Spring, 1480 m, 30.V.1989, A. Smetana (16, CNC); Jackson Co.: highway 140, Little Butte Creek, 23.VI.1974, A.&D. Smetana (17, CNC); Klamath County: 13 mi NE Bly, near Deming Creek, 1706 m, 21.VII.1979, J.M. Campbell & J. Schuh (3, CNC); 16 mi NE Bly, Deming Creek Road, 1828, 21.VII.1979, J.M. Campbell & J. Schuh (9, CNC); 6 mi S Fort Klamath, Crooked Creek, 25.VI.1974, A.&D. Smetana (2, CNC); 9 mi NE Bly, Deming Creek, 1500-1760 m, A.&D. Smetana (6, CNC); Bly Mountain, 24.VI.1974, A.&D. Smetana (1, CNC); Gearhart Mountain, 1980-2194 m, 24.VI.1974, A.&D. Smetana (1, CNC); Sevenmile Creek, 1280 m, 20.VII.1979, J. Schuh & J.M. Campbell (2, CNC); Woodriver Springs, Jackson F. Kimball State Park, 25.VI.1974, A.&D. Smetana (15, CNC); same except, 1295 m, 20.VII.1970, JM Campbell (11, CNC); Mare’s Egg Spring, 1280 m, 20.VII.1979, J. Schuh & JM Campbell (1, CNC); same except, 25.VI.1975, A.&D. Smetana (7, CNC); Tecumseh Spring, 1280 m, 20.VII.1979, J. Schuh & J.M. Campbell (2, CNC); Tillamook Co.: 1 mi S Hebo, 28.VII.1979, J.M. & B.A. Campbell (2, CNC); Umatilla Co.: Umatilla National Forest, North Side Sugarloaf Mountain via Daniel Spring, mixed conifer forest, in wet moss at edge of spring/seep, 12.V.2012, A. Newton & M. Thayer (2, FMNH); 12 km NE Tollgate, Blue Mountain Road 63, 1250 m, A. Smetana (12, CNC); Union Co.: Blue Mountains, 9 km NW Elgin, Philips [Gordon?] Creek Road, 950 m, 27.V.1989, A. Smetana (22, CNC); same except 900 m, 25.V.1989 (3, CNC); Blue Mountains, Road 62, Jarboe Creek, 1200 m, 29.V.1989, A. Smetana (1, CNC); Utah: Cache Co.: Logan Canyon, 2 km N Wood Camp, sifting moss, 1706 m, 14.VII.1981, J.M. Campbell (36, CNC); Washington: Clallam Co.: 5 mi W Forks, 14.V.1968, Campbell and Smetana (1, CNC); Okanogan Co.: 8 mi NNW Republic, Sweat Creek [picnic area], 1097 m, 20.VII.1978, L&N. Herman (1, CNC); Pierce Co.: Mount Rainier National Park, Tahoma Creek, 700 m, 12.VIII.1973, A, Z & D Smetana (14, CNC); same except, 730 m, 10.VIII.1973 (2, CNC); Spokane Co.: 2 mi E Nine Mile, 13.IX.1955, R.A. Ward (1, CNC); Mount Spokane State Park, 1 km NE park entrance, 1000 m, sifting moss, 1.VIII.1981, JM Campbell (19, CNC); Mount Spokane State Park, Bald Knob campground, sifting moss, 1524 m, 31.VII.1981, JM & BA Campbell (1, CNC).

Diagnosis.

Iratiquedius prostans can be distinguished by a combination of elytra with even punctation, not arranged in rows, and pale pubescence at the bases of the abdominal tergites and sternites.

Redescription.

Measurements ♂ (n = 5): HW/HL 1.10-1.14; PW/PL 1.03-1.13; EW/EL 1.23-1.33; ESut/PL 0.65-0.76; PW/HW 1.05-1.15; forebody length 2.5-2.9 mm.

Measurements ♀ (n = 5): HW/HL 1.11-1.16; PW/PL 1.01-1.10; EW/EL 1.23-1.31; ESut/PL 0.69-0.78; PW/HW 1.06-1.09; forebody length 2.8-3.3 mm.

Head dark brown, pronotum and often elytra paler, dark reddish brown to reddish brown, abdominal segments broadly paler apically; antennae dark brown, antennomeres 1-3 with pale base; palpi reddish brown with apical segment dark brown; legs yellowish brown, tibia and metacoxae dark brown, tarsi brownish.

Head slightly transverse, appearing orbicular, temples extremely short, following outline of eye to neck; disc of head with moderately sparse microsculpture of transverse waves, becoming vaguely meshed in places, often completely meshed on frons, where it is denser; posterior frontal puncture located at posterior third of eye; interocular punctures absent; labrum short, transverse, forming two lobes; area between anterior frontal punctures with Y-shaped impression; antennomeres 1-4 or 1-5 elongate, 6-10 subquadrate, nine or ten, sometimes weakly transverse, antennomeres generally becoming shorter toward apex of antennae; pronotum roughly shield-shaped, subquadrate to slightly transverse; disc with microsculpture similar to that of head but becoming meshed on anterior angles; elytra appearing moderately to distinctly transverse; disc without microsculpture, evenly, moderately densely punctate, punctures generally closer than one puncture diameter but only sometimes touching, setae pale yellowish, appearing dark in greasy or wet specimens; abdominal tergites III-V, sometimes weakly on VI, with paired median impressions creating a ‘pinched’ appearance (Fig. 3F ); tergites with paired patches of golden setae, one medial and one occupying entire basolateral corner; sternites with basal areas of golden setae; tergites with microsculpture of very fine and dense transverse waves; tergites with punctation varying from moderately dense at base to very sparse at apex.

Male. Sternite VIII with distinct, moderately deep and rounded emargination; tergite X elongate triangular to triangular, with several long marginal setae; sternite IX distinctly dilated at midlength, with long asymmetrical basal part and moderately deep emargination; median lobe in ventral view with one small, short, median tooth, apex truncate (Fig. 6L, M ); median lobe in lateral view strongly narrowing to small apical part, apex rounded and with small ventral tooth, apical part projecting ventrad (Fig. 6N ); internal sac with paired sclerites including a pair of slender, curved rod-like sclerites and a pair of broader fang-shaped sclerites (Fig. 6O ); paramere longer than median lobe, elongate spoon-shaped in apical half, basal half markedly broad, apex narrowly rounded, peg setae arranged in single, elongate median field (Fig. 6P ).

Female. Female tergite and sternite VIII with apex truncate to vaguely emarginate. Tergite X roughly pentagonal, with basal margin deeply incised, disc with faint to distinct, narrow median sulcus, apical area with inverted U-shaped darkening, apex slightly projected (Fig. 9C ).

Distribution.

Canada: BC. United States: AZ, CA, CO, ID, MT, NM, NV, OR, UT, WA.

Iratiquedius prostans is the most widespread species of the genus. It occurs along the entire western cordillera, including both sides of the continental divide, and as far south as New Mexico in the east and near the United States border with Mexico, in the west. The species is not yet known from mountainous southern Alberta but is expected there.

Bionomics.

Although this species, like other Iratiquedius , seems to prefer moss, it has also been collected in other types of wet litter and even in rotting hay. This broader tolerance of microhabitats has likely allowed for a much wider distribution, across the drier forested areas of the western cordilleras to reach the eastern side of the continental divide.

Comments.

Specimens from across the distribution range were dissected and no consistent differences were observed in the aedeagus. This species varies enormously in size and in proportion of the body, giving the impression of multiple species. All specimens sequenced for CO1 barcodes, including those from both sides of the continental divide, were found to belong to a single cluster with 1.50% maximum pairwise distance (Fig. 10 ).

Figures

Figure 2. A-H dorsal habitus of A, B Iratiquedius amabilis (Smetana) A male holotype B female non-type C, D I. mutator (Smetana) C male non-type D female holotype E I. prostans (Horn) F I. seriatus (Horn) G Paraquedius puncticeps (Horn) H P. marginicollis sp. nov. Scale bars: 1 mm.

Figure 3. A-D dorsal head A, B showing confluence of inner eye margin and supra-antennal carina (arrow) E, F abdominal tergites A, D Iratiquedius seriatus (Horn) B Quedius (Raphirus) probus (Casey) C, E I. amabilis (Smetana) F I. prostans (Horn). Abbreviations: af = anterior frontal puncture; b = basal puncture; pf = posterior frontal puncture; sa = supra-antennal puncture. Scale bars: 0.1 mm (A, B); 0.5 mm (C-F).

Figure 6. A-G, L-O median lobe of aedeagus A-D, N, O in lateral view E-G, L, M in ventral view O with internal sac everted H-K, P underside of paramere showing peg setae A, B, E, F, H, I Iratiquedius amabilis (Smetana) C, D, G, J, K I. mutator (Smetana) L-P I. prostans (Horn). Scale bars: 0.1 mm.

Figure 9. A-I female tergite X A Iratiquedius amabilis (Smetana) B I. mutator (Smetana) C I. prostans (Horn) D I. seriatus (Horn), evenly convex E I. uncifer sp. nov., discal impression F Paraquedius puncticeps (Horn) G P. marginicollis sp. nov. H-I Quediellus debilis (Horn). Pigmented area on tergite in E not shown for clarity but similar to I. seriatus. Scale bars: 0.1 mm.

Figure 10. Neighbor-joining trees of CO 1 barcode sequences for species of Iratiquedius, Paraquedius, and Quediellus, calculated under the Kimura- 2 model for pairwise distance. White circles indicate macropterous ' debilis' morphotypes of Q. debilis, all other Q. debilis are of brachypterous ' nanulus' morphotypes. Scale bar equivalent to 2 % divergence. Sequence length, including number of ambiguous base pairs (N's), is given at the end of each sample name.

Figure 11. Distributions of A Iratiquedius amabilis (Smetana) (circles), I. mutator (Smetana) (squares) B I. prostans (Horn) (black - specimens examined, white - literature records) C I. seriatus (Horn) (circles), I. uncifer sp. nov. (squares) D Paraquedius puncticeps (Horn) (circles), P. marginicollis sp. nov. (squares) E Quediellus debilis (Horn) (circles) (black - specimens examined, white - literature records).