Leucophora johnsoni ( Stein, 1898 )

Leucophora johnsoni ( Stein, 1898) View in CoL

Figs. 30–34 View FIGURE 30 View FIGURE 31 View FIGURE 32 View FIGURE 33 View FIGURE 34

Hammomyia peullae Malloch, 1934 View in CoL

Hammomyia argentina Malloch, 1934 View in CoL

Hammomyia maculipennis Albuquerque, 1953 View in CoL

Hylemyia johnsoni Stein, 1898: 215 View in CoL , Type-locality: Manayunk, Pennsylvania; Aldrich, 1905:552 (catalogue); Stein, 1911:151 (catalogue); Britton, 1920:198 (catalogue); Johnson, 1925:234 (catalogue); Petch & Maltais, 1932:75.

Ammomyia johnsoni Stein, 1919:152 (catalogue); Stein, 1920: 84 (catalogue).

Hammomyia johnsoni Malloch, 1921:78 (key of identification); Huckett, 1924:13 (key of identification); Hallock & Parker, 1926:16 (catalogue); Johannsen, 1928:837 (catalogue); Wilson, 1932:85 (catalog); Petch & Maltais, 1932:76.

Hammomyia peullae Malloch, 1934:192 View in CoL , figs. 27c (male genitalia), d (sternite 5), Type-locality: Chile, Peulla; Ortiz, 1946:156; Albuquerque, 1953:538 (catalogue).

Hammomyia argentina Malloch, 1934:193 View in CoL , fig. 27e (male genitalia), f (sternite 5), Type-locality: Argentina, Mendoza; Albuquerque, 1953:538 (catalogue)

Hylephila johnsoni Séguy, 1937:137 (catalogue).

Leucophora johnsoni Huckett, 1940:336 View in CoL (catalogue), 340, 341 (key of identification), 343 (morphological notes); Drew, 1963:250 (catalogue); Huckett, 1965a:153 (key of identification), 154 (catalogue); Huckett, 1965b:867 (catalogue); Batra, 1965:375 (biological notes); Knerer & Atwood, 1967:105 (biological notes); Huckett, 1972:221 (catalogue); Huckett, 1974:156 (catalogue); Pont, 1974:7 (catalogue); Huckett, 1987:1107, fig. 42 (ovipositor); Griffiths, 1996:1667 (redescriptions and biological notes), 1668 figs. 1803–1806 (male terminalia), 1672 (Nearctic distribution map).

Hammomyia maculipennis Albuquerque, 1953:535 View in CoL Type-locality: Itaquacetuba, Brazil, figs. 1 (arista), 2 (ala), 3 (hind leg), 4 (male genitália frontal view), 5, 6, 7, 8 (aedeagus), 9 (sternite 5), 10 (female cerci), 11 (spermatheca).

Leucophora argentina Pont, 1974:7 View in CoL (catalogue); Ramírez-Mora et al. 2023:323 (catalogue).

Leucophora maculipennis Pont, 1974:7 View in CoL (catalogue); Gomes, 2019:100 (checklist); Ramírez-Mora et al. 2023:323 (catalogue).

Leucophora peullae Pont, 1974:7 View in CoL (catalogue); Polidori et al. 2015 (biological notes); Ramírez-Mora et al. 2023:323 (catalogue).

Diagnosis. Leucophora johnsoni can be differentiated from its congeners by the following combination of characters: transverse veins r-m and/or dm-cu clouded; male with slender surstylus (in posterior view), pregonite with lateral lobe, distiphallus short without free paraphallic processes, and acrophallus with distal cuticular projections; female with abdominal tergite 6 connected at middle, sternites 6–7 widened on posterior margin, tergite 8 very long.

Material examined. Paratype 1 ♂, 1 ♀ ( BMNH): Hammomyia peullae Chile, Peulla ( BMNH) “Peulla: / 12- 13.xii.1926.”; “S Chile: / Llanquihue Prov. / F.&M. Edwards. / B.M.1927–63”; “ Para- / type”. https://data.nhm. ac.uk/object/5d684c35-8195-4cf8-8cea-171dd3b9c6cf https://data.nhm.ac.uk/object/44424503-6242-4706-87aa- 5b711aaf7071

Other material. 62 ♂, 9 ♀ ( IFML, MACN). ARGENTINA: Buenos Aires, Ensenada, Reserva Punta Lara, Malaise , 4 [m.a.s.l.], -34.79964, -58.00745, 15–21-XII-2016, C. Cazorla col. GoogleMaps 2 ♂, 1 ♀ ( MACN _ En 36346, 39858), same data except, 8–15-II-2017 GoogleMaps , 1 ♂, 1 ♀ ( MACN _ En 39860, 39861); PPE Tornquist , Campamento Base Co. Ventana, Transecta , 500 [m.a.s.l.], -38.055280, -61.978819, 19-II-2016, Mulieri, Olea & Patitucci col. GoogleMaps 3 ♂ ( MACN _ En 36293, 36294, 36295), same data except Est. Meteorológica, Transecta , 520 [m.a.s.l.], 20-II-2016, Olea col. GoogleMaps 1 ♂ ( MACN _ En 36296), same data except, Patitucci col. GoogleMaps 1 ♂ ( MACN _ En 36297), same data except, 26-X-2016, Mulieri & Olea col. GoogleMaps 2 ♂ ( MACN _ En 36307, 36308); Ruta 76, Transecta , 247 [m.a.s.l.], -38.171434, -62.341294, 25-X-2016, Mulieri & Olea col. GoogleMaps 4 ♂ ( MACN _ En 36303, 36304, 36305, 36306); same data except, Mulieri col. GoogleMaps 1 ♂ ( MACN _ En 33571); Part. Puan, Barraondo , Transecta , 110 [m.a.s.l.], -38.398225, -62.837612, 29-X-2016, Mulieri col. GoogleMaps 3 ♂, 2 ♀ ( MACN _ En 36318, 36319, 36328, 36329, 36330), same data except, Olea col. GoogleMaps 6 ♂ ( MACN _ En 36320, 36321, 36322, 36323, 36324, 36325), same data except, Pan Trap, Compagnucci col. GoogleMaps 2 ♂ ( MACN _ En 36326, 36327); Pdo Tornquist, Ao. de la Ventana, Transecta , 345 [m.a.s.l.], -38.47894, -61.128122, 19-II-2016, Mulieri col. GoogleMaps 2 ♂ ( MACN _ En 36291, 36292); Ruta Provincial 76, Ao. Atravezado, Transecta , 325 [m.a.s.l.], -38.021876, -61.858028, 21-II-2016, Olea col. GoogleMaps 3 ♂, 1 ♀ ( MACN _ En 36298, 36299, 36301, 36302), same data except, Patitucci col. GoogleMaps 1 ♀ ( MACN _ En 36300), same data except, 30-X-2016, Olea col. GoogleMaps 15 ♂, 2 ♀ ( MACN _ En 33576, 33577, 36331, 36332, 36333, 36334, 36335, 36336, 36337, 36338, 36339, 36340, 36341, 36342, 36343, 36344, 36345); Saldungaray, Río Rivera , Transecta , 246 [m.a.s.l.], -38.231457, -61.800128, 28-X-2016, Mulieri col. GoogleMaps 3 ♂ ( MACN _ En 36309, 36310, 36311), same data except, Olea col. GoogleMaps 6 ♂ ( MACN _ En 36312, 36313, 36314, 36315, 36316, 36317); Villa Ventana , Ao. Belisario, 360 [m.a.s.l.], -38.09070045, -61.925167, 19-II-2016, Mulieri col. GoogleMaps 2 ♂ ( MACN _ En 36289, 36290), same data except, Olea col. GoogleMaps 1 ♂ ( MACN _ En 36288). Catamarca, Angastaco [ -25.691273, -66.167807], 7-XII-1968, A. Willink-Stange col. GoogleMaps 1 ♀ ( IFML). Chubut, PN Los Alerces, Puerto Mermoud , 521 [m.a.s.l.], -42.723190, -71.748825, 25-X-2014, Mulieri col. GoogleMaps 5 ♂ ( MACN _ En 36247, 36248, 36249, 36250, 36251). Salta, Cafayate [ 1625 m.a.s.l.] [ -26.073080, -65.976052], II-1951, Hayward col. GoogleMaps 1 ♀ ( IFML - DIP0890 ). Tucumán, Aconquija [ - 27.327285, -65.771577], 6–10-XII-1950, R. Goldbach col. GoogleMaps ( IFML - DIP0892 ) .

Description

Male ( Figs. 30–32 View FIGURE 30 View FIGURE 31 View FIGURE 32 )

Body length 4.1–6.1 mm, wing length 3.1–4.5 mm.

Coloration ( Fig. 30 View FIGURE 30 ). Head dark brown, with light brown-silvery pruinosity; frontal vitta black to dark brown anteriorly ( Fig. 30B View FIGURE 30 ). Fronto-orbital plate, parafacialia, genae, and face with silvery pruinosity; parafacialia in profile with a wide shifting black to yellow patch from level of insertion of antennae to mid-length of postpedicel ( Fig. 30C View FIGURE 30 ); genae with a brown to yellow patch from the genal margin to the lower margin of the eye. Antennae black, pedicel with dark brown dorsal tip; arista dark brown with black base. Prementum dark brown. Palpi brown on basal half and dark brown distally. Thorax black with greyish and light-brown pruinosity; with three dark brown longitudinal vittae ( Fig. 30D View FIGURE 30 ), central vitta filling the space between the acrostichal setae and not extending over the scutellum (in some specimens from Chubut not extending beyond the first dorsocentral setae), lateral vittae filling the space between the posthumeral and supra-alar presutural setae and intralar postsutural setae and prealar and supra-alar postsutural setae. In some specimens, bases of dorsocentral setae with dark brown base and with light brown pruinosity between the central and lateral vittae; bases of notopleural setae with brown pruinosity. Anterolateral corners of scutellum dark brown. Anepisternum with a yellowish patch ( Fig. 30A View FIGURE 30 ). Legs dark brown; coxae, trochanters, and tibiae lighter; coxae and femora densely covered with greyish pruinosity. Tarsomere almost black. Pulvilli light brown. Wing membrane hyaline with a light brown tinge basally; membrane darkened on r-m and dm-cu transverse veins, in some specimens the darker membrane is not as evident or only visible in one of the transverse veins; veins brown to dark brown. Calypters whitish with a slightly darkened margin. Halteres brown with darker base. Abdomen black with greyish pruinosity; with a central longitudinal dark brown vitta ( Fig. 30E View FIGURE 30 ), narrowed anteriorly on each segment; posterior margin of tergites 3–5 yellowish. Tergites 2–5 with lateral dark brown patch towards posterior of each tergite. Syntergosternite 7+8, epandrium and cerci dark brown. Surstylus and sternite 5 brown, sternite processes lighter.

Head. Fronto-orbital plate at narrowest point 0.8–1.5x width of anterior ocellus, widening anteriorly, bare; frontal vitta at narrowest point 0.8–2.0x width of anterior ocellus, slightly widening towards lunule; eyes separated at narrowest point 2.7–4.6x width of anterior ocellus ( Fig. 30B View FIGURE 30 ). 4–5 pairs of fr s; in some specimens, the uppermost pair short and inserted slightly before the level the anterior ocellus, with setulae in between ( Fig. 30C View FIGURE 30 ). infr s almost as long as the shortest fr s. oc s as long as the longest fr s; ocellar triangle with 2–3 additional pairs of setulae; poc s shorter than the oc s. i vt s slightly shorter than oc s, reclinate; o vt s less than half the i vt s length. Postpedicel 1.2–1.5x the pedicel length; arista 1.9–2.2x the postpedicel length, short plumose, longest hairs 0.5x the postpedicel width. Parafacials 0.3–0.4 width of eye. Genae below lowest point of eye margin 0.2–0.3x eye height. Vibrissal angle slightly projected beyond the frontal angle ( Fig. 30C View FIGURE 30 ). Prementum 0.5–0.6x head height. Palpus 0.4–0.5x as long as head height, slightly expanded at tip.

Thorax. Chaetotaxy: 5 pairs of psut acr s; prescutelar pair longer than the longest presut acr s ( Fig. 30D View FIGURE 30 ). 5 anepst s. 1–2+2 kepst s, lower anterior barely (in some specimens this seta is indistinguishable from the surrounding setulae) ( Fig. 30A View FIGURE 30 ). Prealar seta absent or shorter than anterior notopleural

Legs. Fore femur with a row of av setulae on basal third. Mid femur with 3 short av setae on basal fourth; 4–5 pv setae on basal half; a row of a setae on basal half and 1 subapical seta; 2 p setae. Mid tibia with 1 ad and pd setae; 2 p setae. Hind femur with a row of long av setae on distal half; a complete row of ad setae; 1–2 d setae; 1 p seta; 4 pv setae on basal half and 1 subapical seta. Hind tibia with 3–4 av setae; 3 ad setae; 4 pd setae; 2–3 p setae.

Abdomen. Lateral marginal setae strong ( Fig. 30E View FIGURE 30 ). Sternite 1 about 3x wider than long, setulose except middle, longest setulae almost as long as the sternite width. Sternite 2 slightly longer than wide, narrowed posteriorly. Sternite 3 about 1.4x longer than wide, anterior margin narrowed, posterior margin straight. Sternite 4 slightly longer than wide, shape as sternite 3. Sternites 2–4 setulose, longest setulae on lateral margins of posterior half. Sternite 5 ( Fig. 31A View FIGURE 31 ) about as long as wide, anteriorly tapered, anterior margin almost rounded; processes longer than basal region, slender, apex rounded and narrow, slightly directed inwards, basal inner margin with dense short setulae, outer surface with some short and long setae, longest setae about as long as processes length.

Terminalia . Surstylus longer than epandrium, inner basal margin slightly concave, tips slightly divergent and tapered ( Fig. 31B View FIGURE 31 ); in profile basally convex on dorsal surface, scarcely sinuous at tip, narrowed towards distal half ( Fig. 31C View FIGURE 31 ); some setulae on basal half towards outer lateral surface, otherwise with short sensilla, ventral surface with a longitudinal fringe of setulae on middle. Cercal plate slightly longer than wide, about 0.5x the surstylus length, heart-shape, with rounded apex; in profile with dorsal surface curved at middle, basal and distal apexes narrow and acute; surface setulose and covered with long microtrichia, except on central longitudinal stripe with 4 pairs of long setae, 2 pairs apically and other 2 pairs basally on lateral margin. Pregonite ( Figs. 31E View FIGURE 31 ; 32A View FIGURE 32 ) as long as wide, as long as pregonite, with an outer lateral flat lobe; with 2 setae, 1 preapical seta on the lobe, and 1 seta basally, without sensilla. Postgonite about 2x longer than wide, tapered and pointed distally, seta/setulae absent, only with a few sensilla on basal half on outer surface. Phallapodeme ( Figs. 31D View FIGURE 31 ; 32A View FIGURE 32 ) in profile basally acute and slightly curved, distally sinuous and expanded, unpigmented on apex; epiphallus wide and with rounded apex; distiphallus ( Figs. 31D View FIGURE 31 ; 32A View FIGURE 32 ) short and curved dorsally, without free paraphallic processes; acrophallus cylindrical, with irregular acute projections on distal margin ( Fig. 32B View FIGURE 32 ), unpigmented.

Female ( Figs. 33–34 View FIGURE 33 View FIGURE 34 )

Body length 3.8–5.0 mm, wing length 3.1–3.6 mm.

Differs from male as follows:

Coloration ( Fig. 33 View FIGURE 33 ). Head dark brown to brown, with light grey pruinosity. Ocellar triangle with yellowishsilvery pruinosity ( Figs. 33B, D View FIGURE 33 ). Patches on parafacialia and genae yellowish ( Fig. 33C View FIGURE 33 ). Thorax. In some specimens, with central vitta narrower and the lateral vittae scarcely visible ( Fig. 33D View FIGURE 33 ). Darkened cross-veins of wing more notorious. Abdomen, lateral dark brown patch on tergite 5 may be visible or not ( Fig. 33A View FIGURE 33 ).

Head. Fronto-orbital plate at narrowest point 1.6–2.3x width of anterior ocellus; frontal vitta at narrowest point 1.2–2.2x width of anterior ocellus; eyes separated at narrowest point 4.7–6.4x width of anterior ocellus ( Fig. 33B View FIGURE 33 ). Arista with longest hairs 0.4–0.6x the postpedicel width.

Thorax. 1+1 kepst s, katepisternum with short and scattered setulae ( Fig. 33A View FIGURE 33 ).

Legs. Fore tibia, ad setae stronger. Mid femur with av setae absents; 2 pv setae on basal third. Hind femur with 2 pv setae on basal half and 1 subapical seta.

Abdomen. Lateral marginal setae stronger than male ( Figs. 33A, E View FIGURE 33 ). Sternite 1 slightly more than 2x wider than long, with a couple of setulae on posterolateral corners. Sternite 2 2x longer than wide, slightly narrowed towards posterior margin. Sternites 3–5 1.2x–1.4x longer than wide. Sternites 2–5 setulose, with a pair of long setae on posterolateral margin of each segment.

Ovipositor ( Fig. 34 View FIGURE 34 ). Tergite 6 represented by a wide plate ( Fig. 34A View FIGURE 34 ), slightly wider than long, anterolateral corners narrowly projected anteriorly, with rounded apex; posterior half with a deep but narrow invagination at middle; posterior margin with a row of strong and long setae which extends onto the lateroventral membranes forming with setulae of the sternite 6 a more or less continuous band around segment (longest setae slightly shorter than tergite length). Spiracles 6th and 7th both situated in membrane ( Fig. 34B View FIGURE 34 ), 6th slightly before mid-length of tergite 6, 7th situated in line with setulae in membrane. Tergite 7 represented by two widely divided lateral plates, slightly shorter than tergite 6, straight, not curved; in profile ( Fig. 34C View FIGURE 34 ), anterior margin very height and almost straight, posterior margin rounded; posterior margin with a row of setae of different sizes which extends onto the lateroventral membrane, forming with the setulae of sternite 7 a more or less continuous band around the segment, dorsally the membrane between the lateral plates bare (longest setae shorter than tergite length). Tergite 8 widely divided into a pair of strongly sclerotized lateral plates continuous along posterior margin, the longest tergite, anterior margin strongly tapered; in profile tapered towards anterior margin, apex curved and truncate, posterior margin with rows of very short setulae. When fully extended, tergites 6–8 and sternite 7 remain partially hidden in the anterior segment ( Figs. 34A–C View FIGURE 34 ). Epiproct laterally fused with cerci, slightly more than 2x wider than long, posterior margin narrow and slightly notched, with a pair of long setae. Sternite 6 1.1x wider than long ( Fig. 34B View FIGURE 34 ), shorter than tergite 6, tapered and more sclerotized towards anterior margin, posterior margin with rows of setae. Sternite 7 about 1.2x longer than wider, slightly shorter than sternite 6, anterior margin very narrow with rounded apex, posterior margin less sclerotized and pigmented with rows of setulae. Sternite 8 represented by a pair of small, scarcely sclerotized plates, setulose. Hypoproct wider than long, posterior margin rounded, setulose and pilose. Cerci broad, slightly longer than hypoproct, dorsally with numerous recurved spines ( Fig. 34A View FIGURE 34 ), the longest and stoutest pairs on inner posterior surface (longer than cerci height) ( Fig. 34C View FIGURE 34 ). Spermathecae ( Fig. 34D View FIGURE 34 ).

Remarks. The lobed pregonite in L. johnsoni is shared with L. longilingua and L. yacochuya sp. nov. However, it differs from both species by the presence of the darkened r-m and/or dm-cu transverse veins, also by relatively short distiphallus; from L. longilingua can be separated by the presence of 2 setae on the pregonite, by the short prementum, and the cerci shape; from L. yacochuya sp. nov., can be differentiated by the cerci shape and the absence of the free paraphallic process. Female ovipositor of L. johnsoni resembles that of L. plumiseta in having the tergite 8 enlarged and anteriorly acute, and by the pairs of spiracles 6th and 7th on membrane of segment 6, but differs by the shape of tergites 6–7 and by the wider sternites 6–7 in L. johnsoni .

According to the redescription made by Griffiths (1996:1669), the females of this species have the tergite 6 divided into pairs of dorsolateral plates; however, in the females dissected in this work, the tergite 6 is a wide plate with a deep excavation on posterodorsal margin ( Fig. 36A View FIGURE 36 ).

Griffiths (1996:1667) synonymized L. argentina , L. maculipennis , and L. peullae with L. johnsoni . The three species synonymized were referred to as different species by external characters (the length of the aristal pubescence and the degree of darkening of the transverse veins). This range of variation was observed by Griffiths in the Nearctic material, also this was confirmed by the examination of male genitalia from type material of L. argentina and L. peullae made by Ackland and with the detailed original description and illustration of L. maculipennis . In this work, we previously identified the species L. argentina , L. peullae , and L. maculipennis according to the keys of Malloch (1934) as well as the original descriptions in Malloch (1943) and Albuquerque (1953). We dissected male and female specimens (when possible) from several localities of these species and confirmed the observations made by Griffiths.

Biology. In North America, Batra (1965) found association of L. johnsoni with Lasioglossum zephyrus (Smith) ( Halictidae ) and the kleptoparasitic bee Sphecodes sp. ( Halictidae ); Knerer & Atwood (1967) reported this species as infesting colonies of halictid bees ( Halictus rubicundus (Christ) and Dialectus lineatulus (Crawford). In South America, Polidori et al. (2015) reported for the first time from the Neotropics any anthomyiid fly associated with augochlorine bees. In Chilean Patagonia, they found specimens of L. johnsoni (as L. peullae in the paper) as nest parasites of a new species of Corynura ( Halictidae ), posteriorly described as Corynura ( Corynura) moscosensis González-Vaquero, 2017 .

Distribution. Nearctic and Neotropical ( ARGENTINA, BOLIVIA, BRAZIL, CHILE) ( Fig. 55 View FIGURE 55 ).