Empis (Enoplempis) delumbis Melander

Empis (Enoplempis) delumbis Melander View in CoL

( Figs 143–147 View FIGURES 143–147 , 150, 151 View FIGURES 148–151 , 153 View FIGURES 152–153 )

Empis clauda Coquillett, 1900: 407 View in CoL . Type locality: Popof Island , Alaska ( USA), by lectotype designation.

Empis clauda View in CoL : Melander, 1902: 321 (redescription).

E. (Enoplempis) clauda View in CoL : Melander, 1928: 142 (assigned to subgenus Enoplempis ).

Empis delumbis Melander, 1965: 459 View in CoL (nom. nov. clauda Coquillett View in CoL , catalogue); Yang et al., 2007: 133 (catalogue).

Type material examined. LECTOTYPE (here designated in order to fix identity of the species) ♂, labelled: “Popof Island [55°18′N 160°22′W]/ Alaska, July 12’99”; “Harriman Expedition ’99/ T. Kincaid, Collector”; “ TYPE / No./ 5209/ U.S.N.M. [red label]”; “ Empis / clauda/ Coq. [Coquillett—handwritten]”; “ LECTOTYPE / Empis clauda Coquillett / des. Sinclair, Brooks &/ Cumming 2025 [red label]” [abdomen lost] ( USNM). PARALECTOTYPES: USA. Alaska: Same data as lectotype (1♂, USNM); same data except, 8.vii.1899 (2♂, 1♀, USNM); same data except, 9.vii.1899 (3♀, USNM); same data except, 10.vii.1899 (5♂, 3♀, USNM); same data except, 11.vii.1899 (2♂, 2♀, USNM). Kodiak Island Borough: Kodiak [57°47′N 152°23′W], 20.vii.1899, T. Kincaid (1♀, USNM). Yakutat Borough: Yakutat [59°32′49″N 139°43′38″W], 21.vi.1899, T. Kincaid (1♀, USNM).

Notes on type specimens. The syntype series included 17 males and 14 females from three Alaskan localities: Yakutat, Popof Island and Kadiak [sic] ( Coquillett 1900). In addition, the type catalogue number “5209” was listed for this species. Only one male specimen bears the type catalogue number label and this was selected as the lectotype. Of the original number of specimens in the type series, only 11 male and 11 female syntypes were identified .

Additional material examined. CANADA: British Columbia: Aiyansh, Nass R. [55°12′N 129°4′W], 25. vi.1960, JGC (1♂, CNC); Bowser [49°26′N 124°40′W], 30.v.1955, R. Coyles (1♀, CNC); Vancouver Is., Courtenay [49°41′N 124°59′W], vi.1965, N.L.H. Krauss (1♂, CNC); Exchamsiks R, 35 mi. W Terrace [54°20′N 129°17′W], 9.vii.1960, JGC (1♂, 1♀, CNC); Gagnon Rd, 6 mi. W Terrace [54°29′N 128°44′W], 24.vi.1960, G.E. Shewell (1♀, CNC); same data except, 20.vi. 1960, 220 ft, W.W. Moss (1♂, CNC); Glacier Ck, 10 mi. N Terrace [54°39′N 128°40′W], 7.vii.1960, JGC (2♂, CNC); Graham Is., 1 km W Queen Charlotte City [53°15′N 132°06′W], 26.vii.1981, D.H. Kavanaugh (1♂, CAS); same data except, Queen Charlotte City just east of town, 53°15′10″N 132°02′50″W, 6.viii.2001, S. Allombert, #: GG-01-021 (1♂, RBCM); Haida Gwaii, Graham Is., Bonanza Ck, 4.viii.1983, R.A. Cannings (1♂, RBCM); Haida Gwaii, Gwaii Haanas Pk Res., Gordon Islands, 52°06′N 131°08′W, 5.vii.2000 (1♂, CNC); same data except, Burnaby Is., 52°21′N 131°16′W, 9.vii.2000 (1♂, CNC); same data except, Moresby Is., Garcin Rock, 52°12′N 130°58′W, 7.vii.2000 (1♂, CNC); same data except, Kunghit Is., 52°04′N 131°00′W, 10.vii.2000, grassy area by stream (2♀, CNC); Haida Gwaii, Laskeek Bay, Low Is., 52°54′38″N 131°32′07″W, 24.vii.2001, M. Creery, #: LG-01-035 (1♂, RBCM); same data except, 18.vii.2001, #: LG-01-031 (1♂, RBCM); same data except, 131°32′10″W, 19.vii.2001, S. Allombert #: LI-2SWE-2 (1♀, RBCM); same data except, Reef Is. 52°52′39″N 131°31′07″W, 15.vii.2001, #: RG-01-133 (1♀, RBCM); same data except, West Skedans Is., 52°57′26″N 131°34′42″W, 24.vii.2001, #: WI-3SWE-3 (1♂, RBCM); same data except, South Skedans Is., 52°57′00″N 131°34′00″W, 18.vii.2001, #: SI-1SWE-2 (1♂, RBCM); same data except, Lost Is., 52°48′14″N 131°29′11″W, 25.vi.2001, #: TS-6G02-3 (1♂, RBCM); Kleanza Ck, 14 mi. E Terrace [54°35′N 128°24′W], 17.vi.1960, CHM (5♂, 6♀, CNC); same data except, 29.vi.1960 (11♂, 2♀, CNC); same data except, 30.vi.1960 (14♂, 11♀, CNC); same data except, 4.vii.1960 (3♂, 3♀, CNC); same data except, 13.vii.1960 (1♀, CNC); same data except 17.vi.1960, JGT (1♂, 6♀, CNC); same data except, 29.vi.1960 (3♂, 1♀, CNC); same data except, 30.vi.1960 (4♂, 1♀, CNC); same data except, 4.vii.1960 (3♂, 1♀, CNC); same data except, 13.vii.1960 (3♂, CNC); same data except, W. R. Richards (1♂, 1♀, CNC); same data except, 19.vii.1960 (1♂, 4♀, CNC); same data except, 12.vii.1960, G.P. Holland (1♂, CNC); same data except, 30.vi. 1960, 250 ft, B. Heming (2♀, CNC); same data except, 25.vii.1960 (1♂, 2♀, CNC); same data except, 17.vi.1960, R. Pilfrey (1♂, 2♀, CNC); same data except, W.W. Moss (2♀, CNC); same data except, 14.vii.1960, G.E. Shewell (1♂, CNC); Lakelse, nr Terrace [54°23′N 128°31′W], 300 ft, 14.vi.1960, W.W. Moss (1♂, 1♀, CNC); Lakelse Lk. bog, S of Terrace [54°23′N 128°31′W], 14.vi.1960, JGC (1♂, 2♀, CNC); same data except, B. Heming (1♀, CNC); same data except, 27.vi.1960 (1♂, CNC); Lulu I. [49°10′N 123°04′W], 10.vi.1916 (1♂, 1♀, USNM); Moresby Is., Mosquito Lk. [53°04′N 132°04′W], 20.vii.1988, meadow nr lake, T.A. Wheeler (1♀, CNC); Mt Thornhill nr Terrace [54°30′N 128°35′W], 14.vii.1960, JGC (1♂, 1♀, CNC); same data except, 26.vii.1960, B. Heming (1♂, 1♀, CNC); same data except, 30.vii.1960; Nares Islets [54°29′N 130°50′W], 29.vii.1987, largest islet open forest, C.S. Guppy (2♂, 1♀, RBCM); Prince Leboo Is. [54 27N 130 59W], 1 km E on small island shore & forest, 25.vii.1987, C.S. Guppy (2♂, 1♀, RBCM); Qualicum [49°21′N 124°26′W], 15.vi.1955, G.E. Shewell (1♀, CNC); 13 km SE Port Renfrew [48°30′N 124°17′W], 5.vii.1991, A. Borkent, CD1340 (1♂, CNC); Shames, 18 mi. SW Terrace [54°29′N 128°57′W], 17.vii.1960, CHM (1♀, CNC); same data except, 104 ft, 17.vii.1960, B. Heming (1♂, 1♀, CNC); Spring Ck, Terrace [54°30′N 128°35′W], 11.vi.1960, CHM (2♂, 1♀, CNC); Terrace, 220 ft, 19.vii.1960, B. Heming (3♀, CNC); same data except, Ferry Island, 8.vi.1985, R.A. Cannings (1♀, RBCM); 32 mi. SW Terrace [54°20′N 129°17′W], 9.vii.1960, JGC (1♂, CNC); same data except, 11.vi.1960 (2♀, CNC); same data except, 6.vii.1960, CHM (1♀, CNC); same data except, 100 ft, 5.vi.1960, B. Heming (1♂, 2♀, CNC); same data except, 9.vii.1960 (1♀, CNC); same data except, 13.vii.1960 (2♂, 1♀, CNC); 54 mi. SW Terrace [54°11′N 129°39′W], CHM (1♀, CNC); Zymoetz R, 6 mi. E Terrace [54°32′N 128°28′W], 3.vi.1960, JGC (1♀ CNC). USA. Alaska: Aleutians East Borough: On tundra, Cold Bay, 163°W [55°12′N 162°42′W], 13.viii.1952, W. R. Mason (3♂, 4♀, CNC); same data except, 28.vii.1952 (2♀, CNC); same data except, 18.viii.1952 (2♂, CNC); same data except, 20.viii.1952 (1♂, 1♀, CNC); same data except, 21.viii.1952 (1♂, CNC); same data except, 25.viii.1952 (7♂, 5♀, CNC); same data except, 26.viii.1952 (1♂, 1♀, CNC); same data except, 27.viii.1952 (2♀, CNC); same data except, 28.viii.1952 (1♂, 1♀, CNC); Aleutians West Census Area: Unalaska [53°53′N 166°31′W], 24.vii.1897 (1♂, 4♀, USNM); Unalaska, Makushin [53°54′N 166°42′W], 11.viii.1907, E.C. Van Dyke (2♂, 2♀, CAS). Juneau Borough: Douglas [58°16′N 134°23′W], 25.vii.1901 (1♀, USNM). Kenai Peninsula Borough: Seward, 60.11242°N 149.35723°W, Populus spp. , pruned branch tips, 21.vii.2014, R. Progar, S. Bresney (1♂, UAM). Ketchikan Gateway Borough: Revillagigedo Is. [55°19′N 131°31′W], Herring Cove, 1.vii.1947, Reed (1♀, CAS). Kodiak Island Borough: Kodiak, Near Is. [57°46′N 152°24′W], 0–30 m, 57.78813°N 152.39081°W, Picea rainforest, Heracleum umbels, rocky beach, 29.vi.2013, I. MacDougall (1♀, UAM). Oregon: Clackamas County: Mt Hood NF, Tollgate Cpgd [45°19′N 121°54′W], T 3S R 7E Sec. 11, 22.vi.1977, DDW (4♂, 2♀, CAS). Clatsop County: Saddle Mtn [45°57′N 123°41′W], 14.vi.1969, K. Goeden (1♂, ODAC); S slope Saddle Mtn, 2500–3280 ft, 14.vi.1969, E.M. Fisher (1♂, CSCA). Hood River County: Starvation Ck SP [45°41′N 121°41′W], at falls, 26.vi.1978, DDW (1♀, CAS). Lane County: H.J. Andrews Exp. For. [44°13′N 122°10′W], 1450 ft, R.S.7, 18.v.1983, sweep, D.S. Chandler (3♂, UNHC). Linn County: 0.5 mi. past Tombstone Pass, Hackleman Ck, 44°23′51″N 122°7′53″W, 29.vi.2016, SJF (1♂, CNC); Willamette NF, 1007 m, Hackleman Ck Rd No. 2672, 44°24.733′N 122°02.555′W, 11.vii.2013, BJS (1♀, CNC); Willamette NF, 335 m, Rd 2025 along Moose Ck, 44°25.184′N 122°24.987′W, 11.vii.2013, streams, BJS (1♀, CNC). Tillamook County: Cascade Head Exp. For. [45°03′N 123°57′W], 24.vi.1961, S.L. Wood, J.B. Karren, D.E. Bright (1♂, BYU). Washington: Clallam County: Bogachiel, Bogachiel R [47°53′N 124°21′W], 29.vi.1974, PHA (1♀, CAS); Dean Ck, 7 mi. S Sequim [48°00′N 123°02′W], 4.viii.1972, WJT, W.B. Garrett (1♂, WSU); same data except, Dry Ice MT (1♂, WSU); Olympic NP, Deer Pk, Blue Mt. [47°57′N 123°15′W], 1650 m, 23.vii.1978, DDW (1♂, CAS); Olympic NP, Elwha Valley entrance (E side Lk. Mills) [48°01′N 123°35′W], 20.vii.1983, 1585/1/6, J.A. Downes (1♂, CNC); Olympic NP, Heart of the Hills [48°05′N 123°25′W], 15.vii.1977, R.S. & V.L. Zack (1♀, WSU); Olympic NP, Lk. Crescent, waterfalls W Storm King Visitor Center [48°03′N 123°47′W], 30.vi.1974, PHA (1♀, CAS); Olympic NP, along Lake Ck [48°01′N 123°24′W], 650 m, 24.vi.1978, DDW (1♀, CAS); Olympic NP, 11 km SSE Solduc Cpgd, entrance to loop B, Hemlock for., sweep nr fallen logs, 47°57′55″N 123°51′11″W, 545 m, 18.vii.2005, C.J. Borkent, CJB05-32 (1♂, LEM). Grays Harbor County: Preachers Slough E., Chehalis R Surge Plain Preserve, 46°56′41″N 123°39′01″W, 16.vii.2024, J.M. Cumming (2♀, CNC). Jefferson County: 17 mi. S Forks, Cottonwood Cpgd on Hoh R [47.7788°N 124.2930°W], 17–18.vii. 1977, 200 ft, WJT (1♀, WSU); Hoh R Cons. & Rec. Corridor, riparian rainforest, 47°49′14″N 124°12′09″W, 20.vii.2024, J.M. Cumming (3♀, CNC); falls outside Olympic NP entrance, 15 km NE Quinault [47°31′N 123°42′W], 28.vi.1974, PHA (1♀, CAS); 2 mi. NW Triton, UFS Rt. 251 [47°36′N 122°59′W], 3.viii.1972, MT dry ice, WJT, W.B Garnett (1♂, WSU); Quilcene [47°49′N 122°53′W], 24.vii.1917, ALM (4♂, 2♀, USNM). King County: Seattle [47°36′N 122°19′W], 19.vi.1920, E.C. Van Dyke (2♂, CAS); same data except, 16.vi.1920, ALM (1♂, 1♀, USNM); 8 mi. W Snoqualmie Pass [47°23′N 121°28′W], 29.vi.1924, ALM (1♂, USNM). Kittitas County: Lk. Keechelus [47.35°N 121.37°W], 29.vi.1924, ALM (1♂, USNM). Lewis County: Big Ck, 2030 ft, 46°43′50″N 121°54′49″W, 15.vi.2018, SEB (1♂, 1♀, CNC); same data except, 16.vi.2018, J.M. Cumming (1♀, CNC); Big Ck Cpgd [46°44′N 121°58′W], 4.4 km SE Ashford, 2.vii. 1974, 550 m, PHA (1♂, 1♀, CAS). Mason County: Lk. Cushman [47°25′N 123°13′W], 27.vi.1917, H.G. Dyar (4♂, 2♀, USNM); Lilliwaup [47°27′N 123°06′W], 23.vii.1917, ALM (1♀, USNM); Shelton [47°12′N 123°6′W], Walkers Pk, 21.vii.1917, ALM (1♀, USNM). Pacific County: Fort Canby SP [46°17′N 124°03′W], nr Ilwaco, 19.vii.1978, DDW (1♂ with Agromyzidae prey, 7♂, 4♀, CAS); same data except, 13– 15.vi.1973, WJT (1♂, 1♀, WFBM); Ilwaco [46°18′N 124°01′W], viii.1917, ALM (1♂, 2♀, USNM); same data except, 28.vi.1925 (3♂, USNM); same data except, viii.1918, E. Miner (2♀, USNM); Naselle [46°22′N 123°47′W], 2.vii.1925, ALM (1♂, 1♀, USNM); Naselle R, 15.vi.1922 (1♂, USNM). Pierce County: Mt. Rainier NP, Fish Ck at West Side Rd [46°47′N 121°53′W], 910 m, 1.viii.1977, DDW (1♂, 1♀, CAS); Mt Rainier, Base No. Side [46°57′N 121°47′W], 26.vii.1924, ALM (1♂, USNM); Mt Rainier, White R [46°54′N 121°38′W], 20.vii.1924, ALM (1♀, USNM); Mt Rainier, Tahoma Fork [46°44′N 121°53′W], 25.vii.1922, ALM (1♂, USNM); Mt Rainier, Longmire [46°45′N 121°48′W], 20.vii.1922 ALM (1♂, USNM); Mt Rainier, Cougar Rock to Longmire Tr., 46°45.6′N 121°48.2′W, 17.vii. 2012, 920 m, BJS (1♂, 1♀, CNC). San Juan County: Friday Harbor [48°32′N 123°01′W], 1.vi.1906 (1♀, USNM); Mt Constitution [48°40′N 122°49′W], 22.vii.1909 (3♂, 1♀, USNM). Skagit County: Anacortes [48°30′N 122°37′W], W. Mann (3♂, USNM); Anacortes, 28.vi.1909 (3♂, USNM); Mt Vernon [48°25′N 122°19′W], 3.vii.1924, ALM (7♂, USNM). Skamania County: 3 mi. W Spirit Lk. Cpgd, Rt. 504 [46°17′N 122°11′W], 30.vii.1972, MT with dry ice, WJT,W.B. Garnett (1♂, WSU). Snohomish County: Big Four Mtn [48°02′N 121°31′W], 5.vii.1924, ALM (18♂, 6♀, USNM); Everett [47°58′N 122°12′W], 19.vi.1920, ALM (10♂, 3♀, USNM); same data except, 4.vii.1924 (1♂, USNM). Whatcom County: Middle Fk Nooksack R, 900 ft, 48°46′15″N 122°4′27″W, 11.vi.2018, nr dam, S.H. Cumming (1♀, CNC). Yakima County: 10 mi. E Chinook Pass summit [46°56′N 121°21′W], 31.vii.1976, N.E. Woodley (1♂, USNM).

Diagnosis. This species is distinguished from other species of the E. (En.) delumbis species group by the yellow colouration (occasionally thorax brown), male hind tibia bearing anteroventral lobe without setae, slightly arched with flattened anterior face ( Figs 146, 147 View FIGURES 143–147 ), phallus with subapical expansion with spiculate surface and pair of narrow lateral lobes.

Redescription. Wing length 6.0– 7 mm. Male. Head dark in ground-colour, with greyish pruinescence. Eyes closely approximated, upper ommatidia not enlarged. Frons very narrow, divergent towards antennae; below ocellar triangle narrower than width of anterior ocellus; with short setulae along inner margin of eye. Face without setae; slightly divergent towards mouthparts, with oral margin dark and shiny. Ocellar triangle elevated, with thin greyish pruinescence, with pair of parallel ocellar setae, shorter than pprn, and several pairs of short setulae. Occiput bearing single row of strong setae, similar to vertical setae, longer and finer ventrally; postocular setae shorter and finer than occipital setae; postocellar region without setae. Antenna brown; scape 2× longer than pedicel, with short, black setae; pedicel with ring of apical setae; postpedicel long and tapered, 4× longer than basal width; stylus 2× longer than basal width of postpedicel. Palpus yellow, with dark setulae. Proboscis reddish brown, length 2× or less head height; labellum with short, dark setae.

Thorax mostly yellow in ground-colour ( Fig. 144 View FIGURES 143–147 ), occasionally brown ( Fig. 143 View FIGURES 143–147 ), densely whitish pruinescent. Scutum colouration variable (dependent on preservation method), ranging from mostly yellow with broad brownish vittae between acr and dc rows and darker outer vittae to mostly dark or rarely with narrow dark vittae beneath dc; postpronotal lobe, lateral margin of scutum, postalar ridge and scutellum yellowish. Pleura yellow, lower half of katepisternum, anepisternum, meron and metakatepisternum often dark brown; mediotergite sometimes with pair of large, dark ventral spots. Proepisternum with short, dark setae on lower section; upper proepisternum in front of spiracle without setae. Prosternum bare. Antepronotum with row of dark setae. Postpronotum with 1 long seta and several dark setae of various thicknesses. Scutum with short, fine biserial acr; dc slightly longer than acr, with 1 prescutellar seta distinctly longer and stouter; 1–2 posterior npl, with several short anterior npl setulae; 1 presut spal; 1 psut spal; 1 pal. Scutellum with 2 pairs of sctl, outer pair shorter; disc bare. Laterotergite with patch of strong, dark setae. Anterior and posterior spiracles pale.

Legs slender, yellow, with apical tarsomeres darker; coxae clothed in whitish pruinescence, concolourous with pleura ( Figs 143, 144 View FIGURES 143–147 ). Coxae with short black setae. Hind trochanter with short setulae. Fore femur clothed in short, black setae. Fore tibia with 3–5 erect anterodorsal setae, longer than width of tibia; apex with circlet of preapical setae; anterior face with short appressed setae. Fore tarsomere 1 slightly inflated, wider than apical tarsomeres, with 1 long dorsal seta, more than twice width of tarsomere, with fine ventral setae and biserial row of stronger setae, longer than width of tarsomere; tarsomeres 2–5 with biserial row of spine-like ventral setae; tarsomeres 1–4 with apicolateral pair of spine-like setae, longer than width of segment. Mid femur similar to fore femur, with strong preapical anterior seta. Mid tibia with 3–4 posteroventral setae; 1 long, ventral preapical seta slightly longer than apical width of femur; 4–5 erect anterodorsal and posterodorsal setae; posterior face with dense fine setulae and several erect, strong setae; circlet of strong preapical setae. Mid tarsus with long, strong ventral setae on tarsomere 1; tarsomeres 2–5 similar to fore tarsus. Hind femur with apex somewhat inflated; anteroventral process knob-like, arched posteriorly with setae on outer face, denser on apex and strong setulae at base; tuft of thickened setae distal to process; circular ridge with thinly sclerotized centre distal to tuft; apex with preapical row of 4–5 flattened, black posterior setae; preapical anterior seta strong, longer than width of femur; 1–2 preapical anterodorsal setae ( Figs 145–147 View FIGURES 143–147 ). Hind tibia with subbasal swelling and bend, bearing anteroventral lobe without setae, slightly arched with flattened anterior face; posteroventral swelling tapered to small ridge, with swelling appressed into depression and circular ridge of femur (when legs folded); with seta in posteroapical comb; long, numerous anterodorsal and posterodorsal setae, length more than twice width of tibia; long setae on anterior and ventral surfaces ( Figs 145–147 View FIGURES 143–147 ). Hind tarsomere 1 slightly inflated with long dorsal setae similar to tibia; remaining tarsomeres similar to fore tarsus.

Wing infuscate with dark veins; veins mostly complete (except Sc), CuA+CuP weakly sclerotized, often fading out before wing margin. Basal costal seta absent. R 5 and M 1 divergent near wing margin; R 5 ending posterior to wing tip; radial fork acute. Halter yellow.

Abdomen variable in colour (dependent on preservation method) yellow, with central vitta to dark, with greyish pruinescence, darker than scutum; with long dark setae on posterior margin. Pregenital segments unmodified, except sclerites of segment 8 narrowly fused anteriorly. Terminalia ( Figs 150, 151 View FIGURES 148–151 ): yellow with ventral margin of epandrium, cerci and hypandrium brown. Cercus short, with triangular anterior half; posterior half digitiform, projecting obliquely from epandrium; setae nearly as long as width of cercus. Subepandrial process extending to either side of phallus. Epandrial lamella triangular, broad, width twice width of cercus; posterior tapered, with short, upturned projection, bearing setae ventrally, length about half width of epandrium. Hypandrium long, sharply tapered, pointed medially; setae longer than width of hypandrium. Phallus ( Fig. 151 View FIGURES 148–151 ) with basal half slightly broader than apical half, strongly bent beyond mid-length, with lateral digitiform processes flanking apical section; apex emerging beyond epandrium; apical portion with subapical expansion with spiculate surface, sometimes spicules extending to apex; apex broad, ventral margin somewhat prolonged; ejaculatory apodeme more than half-length of epandrium, Y-shaped, with ventrally positioned lateral apodemes.

Female. Similar to male, except as follows: frons broader, wider than anterior ocellus; hindleg without modified setae and processes; abdominal pleural membrane without darkened, expanded region; tergites 7 and 8 with pruinescence.

Geographical distribution and seasonal occurrence ( Fig. 153 View FIGURES 152–153 ). Empis (En.) delumbis is known from Alaska, British Columbia, Washington and Oregon. Adults have been collected from June to August.

Nuptial gift presentation. A male specimen was examined with an unwrapped fly ( Agromyzidae ) prey item pinned below the specimen. This may indicate that males of this species present unwrapped prey to females as nuptial gifts.