Liriomyza brassicae (Riley)

Lonsdale, Owen, 2011, The Liriomyza (Agromyzidae: Schizophora: Diptera) of California 2850, Zootaxa 2850 (1), pp. 1-123 : 33-38

Liriomyza brassicae (Riley)

Figs 54–68

Phytomyza diminuta . Nomen dubium . Walker 1858: 233. Syn. Frick (1952b).

Oscinis brassicae Riley 1885: 322 .

Agromyza pascuum Meigen 1830 . Misidentification. Melander 1913: 258; Frick 1952a: 402.

Liriomyza cruciferarum Hering 1927: 461 . Syn. Frick (1952b).

Liriomyza brassicae . Frick 1952a: 402, 1957: 68, 1959: 402; Spencer 1959: 309, 1963a: 332, 1963b: 356, 1969: 170; Spencer & Steyskal 1986: 127; Deeming 2006: 409; Gil-Ortiz et al. 2008: 573, Palacios et al. 2008: 12.

Agromyza diminuta Walker. Misidentification , in part. Coquillett 1898: 78.

Phytomyza mitis Curran 1931: 97 . Frick 1952a: 427, 1959: 402. Syn. Spencer (1967 [?]) [not explicit].

Liriomyza hawaiiensis Frick 1952b: 513 . Syn. Spencer (1963b).

Liriomyza bulnesiae Spencer 1963b: 360 . Syn. Spencer & Stegmaier (1973).

Liriomyza ornephila Garg 1971: 241 . Syn. Sasakawa (1977).

Description. Wing length 1.2–1.6mm (♂), 1.7–1.8mm (♀). Length of ultimate section of vein CuA 1 divided by penultimate section: 2.5–3.5. Eye height divided by gena height: 2.9–5.0. Scutum shining.

Chaetotaxy: Two or three (rare) ori, two ors. Acrostichal setulae in four rows.

Colouration: Calypter margin and hairs grey. Lateral margin of frons sometimes with thin brown margin, varying in strength from indistinct (common) to reaching base of fronto-orbitals (rare); posterolateral corner of frons brown to base of inner or outer vertical bristles, sometimes light brown between base of bristles; remainder of head light yellow with back of head and ocellar triangle brown. Scutum with complete lateral yellow stripe; katatergite yellow with posteroventral margin brown; anatergite light brown lateral to scutellum with dorsum yellow and dark below scutellum. Anepisternum usually with most of ventral margin brown, although sometimes also with posterior margin broadly pigmented or only with small anteroventral spot; anepimeron mottled; meron brown with dorsal 1/3 yellow; katepisternum with large brown triangular spot (not enclosing bristle). Legs yellow with tibiae, tarsi and base of fore coxa brown (lighter on fore and mid legs, particularly towards apex); sometimes base of femora (often only dorsally) and scraper on hind femur brown; uncommonly with brown streaking on fore femur, but if present, then lateral margin of frons thinly brown, dark line present between base of vertical bristles, and hind coxa brown. Abdomen brown with lateral and sometimes posterior margin of tergites yellow; tergite 2 sometimes with yellowish mottling, tergites 2–4 sometimes with thin medial dividing yellow line, and tergite 4 often nearly divided medially into two pairs of connected spots; tergite 5 yellow with large brown medial spot; epandrium with yellow dorsal mottling; some California males with abdomen only faintly brownish dorsally.

Genitalia: Figs 54–60 . Surstylus with prominent apical spine and smaller ventromedial spine; less commonly with spines closer and subequal in length. Hypophallus small, thin, curved anteriorly and with few apical hairs. Paraphallus thin or slightly expanded distally. Distiphallus thin, cylindrical, slightly constricted before midpoint (ie. at point of fusion between distiphallus and mesophallus) and dark with pale apical chamber; distal half sometimes wider, appearing as Lantana -reared material discussed below. Ejaculatory apodeme dark and broad, with corners pronounced.

Variation: “Brassicae 2” ( Figs 61–63 ) collected on Foeniculum vulgare (Umbelliferae) as adult: wing length 1.5mm (♂); length of ultimate section of vein CuA 1 divided by penultimate section 3.3–3.4; eye height divided by gena height 3.8–4.8; first flagellomere dark yellow with basal margin pale; lateral margin of frons very thinly brownish; posterolateral corner of frons to base of outer vertical bristle and thin line in front of inner vertical brown, and space between vertical bristles light brown; katatergite brown; anepisternum with oblique clavate stripe; base of femora and streaking on fore femur brown; scraper brownish; tibiae and tarsi brown; ejaculatory apodeme slightly thinner; surstylus with two closely-spaced subequal subapical spines; distiphallus nearly parallelsided in ventral view (slightly wider subapically) and strongly compressed dorsoventrally on basal half.

“Brassicae 3” ( Figs 64–66 ), host unknown: wing length approximately 1.5mm (♂); length of ultimate section of vein CuA 1 divided by penultimate section indeterminate; eye height divided by gena height 4.5; first flagellomere dark yellow with basal and inner-ventral margins paler; face brown; lateral margin of frons to base of frontoorbitals with faint brownish stripe; posterolateral corner of frons dark to base of outer vertical bristle and brownish to base of inner vertical; dorsal 1/3 of anepisternum yellow with yellowish posterior emargination; katatergite brownish ventrally; coxae brown with apex of fore coxa yellow; base of femora and dorsal mottling brown; tibiae and tarsi brown; surstylus with two subequal subapical spines; paraphallus thin and relatively dark; distiphallus barely constricted medially.

Specimens reared from Lantana (Verbenaceae) ( Figs 67, 68 ) and allied material of unknown hosts, differ as follows: wing length 1.5–1.9mm (♂), 1.9mm (♀); length of ultimate section of vein CuA 1 divided by penultimate section 2.6–3.7; eye height divided by gena height 4.2–4.8; dm-cu sometimes incomplete; lateral margin of frons usually yellow, sometimes faintly brownish laterally; posterolateral corner of frons usually only brown to base of outer vertical bristle but sometimes light brown to base of inner vertical; anepisternum sometimes with ventral 2/3 brown; mid and hind femora sometimes brownish dorsoapically and hind femur always at least thinly brown dorsobasally; tibiae sometimes paler on anterior legs, and sometimes with base and apex of fore tibia yellow; abdomen thinly to broadly yellow laterally; epandrium sometimes entirely brown; paraphallus slightly thinner and darker; distiphallus more strongly constricted medially and distal half wider with apical chamber slightly larger.

Hosts. See table 1. Adult collected on Foeniculum vulgare (Umbelliferae) .

Range. Widespread in Nearctic, Neotropical, Oriental, Afrotropical and Australasian Regions. Europe. Arabian Peninsula. Japan.

Type material. Oscinis brassicae : Holotype, USA. Montana: St. Louis, 30.iv.1876 (1♀, USNM). Phytomyza mitis : Holotype, CANADA. Manitoba: Aweme, 20.vii.1929, R.H. Handford (1♀, CNC); Paratypes examined, Manitoba: Aweme, 3.ix.1929, R.H. Handford (2♂, CNC). Liriomyza hawaiiensis : Holotype, USA. Hawaii: Oahu, Honolulu, 1.i.1947, E.C. Zimmerman, ex. leaf of Cleome (1♀, BPBM) [Not examined]. Liriomyza bulnesiae : Holotype, VENEZUELA. Caracas, Botanical Gardens, caught on Bulnesia arborea Engl. (Zygophyllaceae) , 5.xii.1958, K.A. Spencer (1♂, BMNH) [Not examined]. Liriomyza ornephila : Holotype, INDIA. Uttar Pradesh, Pithoragarh (1♀, depository not given) [Not examined]. Liriomyza cruciferarum : Syntypes, CANARY ISLANDS. La Palma: Santa Cruz (2?, ZMHU) [Not examined].

Additional material examined. USA. California: Alameda Co., Berkeley, “Aug. 23/17” (1♂ 1♀, EMEC) , Berkley, “57”, on Brassica nigra, W.W. Jones (2♀, EMEC) , Berkeley , 17.iii.1954, E. Oatman, Columbine, coll. 3.iii.1954 (1♂, EMEC) , Contra Costa Co., Moraga , 21.v.1977, D.G. Denning (1♂, UCD) , El Dorado Co., Riverton , 19.viii.1953, E.I. Schlinger (1♂, UCD) , Snowline Camp , 2.viii.1943, ex. larva Brassica arvensis L., lot No. 70-1, K.E. Frick (1♂ 1♀, EMEC) , 2mi N Kelsey , 23.vi.1967, W.J. Turner (1♂, EMEC) , Imperial Co., S End Chocolate Mts., Ogilby Rd. , 3mi S Jct. Hwy 78, malaise trap, 7AM-5 PM, 20–22.iii.1978, Wasbauer, Slansky & Adams (1♂, CSCA) , Algodones Dunes, Niland-Glamis Rd., 2.6km n. Hwy 78, 33°00’N, 115°06’W, 7–9.ii.2008, S.L. Heydon & T. J. Zavortink, 80m (1♂, UCD) , 85m, on sand dunes (1♂, UCD) , 85m, wash below dunes, Malaise (3♂, UCD) , Algodones Dunes, Wash Road , 10.4km SE Glamis, 32°55.4’N, 114°59’W, 29.iv–2.v.2008, S.L. Heydon & K. Lorenzen, microphyll forest, Malaise (1♂, UCD) , Algodones Dunes, Buttercup Region, 1.6km ESE Gordon’s Well exit & Hwy. 8, 32°45.5’N, 114°57.5’W, 24–28.iii.2008, Bohart Museum Survey Team, MT in Creosote (1♂, UCD) , Inyo Co., Mt. Whit. F. Hatchery , 5mi NW Independence, 28–29.viii.1979, M. Wasbauer & P. Adams, Malaise trap, 8 AM –6 PM (1♂ 2♀, CSCA) , Los Angeles Co., Baldwin Park , 20.viii.1945, J.C. Elmore, “TC639cauliflower” (4♂ 23♀, USNM) , Gardena , 7.vi.1948, ex. Cauliflower (1?, UCD) , Glendale , 30.vii.1955, E.I. Schlinger (1♂, UCD) , Marin Co., Bolinas , 2mi, 5.x.1968, P.A. Rude (1♂, EMEC) , Mendocino Co., Ukiah , 8.x.1964, sweeping ground cherry (2♂, UCD) , Modoc Co., Willow Rch. , 3mi E, 6.vi.1970, Isatis tinctoria, P. Opler (1♂, EMEC) , Orange Co., Long Beach , 26.v.1951, W.W. Jones (1♂, USNM) , El Toro, E. R. Oatman , collected on cabbage, 1.xii.1964 (1♂, UCR) , 13.iv.1963 (1♂, UCR) , Riverside Co., Riverside, A.L. Melander , 22.ii.1935 (1♂, USNM) , Univ. Cal. River campus, 18.x.1978, E.M. Fisher (2♂, CASC) , Fullerton , 13.ii.1969, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♂, UCR) , Yorba Linda , 30.xii.1968, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♀, UCR) , India , 25.iii.1969, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♀, UCR) , Riverside, malaise, 20.vi,1971, M.E. Irwin (1♂, UCR) , San Bernardino Co., New York Mts., 5400’, Keystone Cyn. , 4.5mi S Ivanpah, 12.iv.1978, Malaise trap, 8 AM –5 PM, M. Wasbauer & T. Eichlin (1♂, CSCA) , San Diego Co., Borrego-Clark L. N. End , 23.iii.1978, Wasbauer, Slansky & Adams, Malaise trap, 8 AM –5 PM (2♂, CSCA) , Escondido, E. R. Oatman , 9.vii.1964 (1♂, UCR) , 20.viii.1964 (1♂, UCR) , Jamul , 5.vi.1969, insectary reared on Ambrosia psilostachys Decandolle (1♂, UCR) , Alpine , 28.ii.1970, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♀, UCR) , San Onofre Bluff , 33°21’38”N, 117°32’22”W, 4.ix.1999, Yanega & Gates (1♂, UCR) , Santa Barbara Co., Sta. Cruz. Isl., Water Cyn. , 400’, 17.vi.1967, R.O. Schuster (1♂, UCD) , Sta. Cruz Isl., Christi Beach, 29.iv.1969, D.S. Horning, Jr. (1♀, UCD), UC Coal Oil Pt. Reserve , 34.4105’N, 119.8798’W, Malaise , M. Caterino & A. Borrell, 8–22.iv.2003 (3♂, SBMN) , Los Prietos , 23.vi.1965, J. Powell (1♂, EMEC) , Santa Barbara , 22.iv.1969, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♀, UCR) , Santa Clara Co., Stanford U., P.H. Arnaud , Jr. , 1.v.1961 (1♂, CASC) , Stanford Univ. , 9.x.1950, P.H. Arnaud, Jr. (1♂, USNM) , San Jose , 15.x.1948, P.H. Arnaud, Jr. (1♂, USNM) , Vacaville , 8.vi.1953, E. Mexger (1♂, UCD) , Sonoma Co., Healdsburg , 8.x.1964, sweeping Dandelion (1♂ 1♀, UCD) , Ventura Co., beach nr. Punta Gorda , 27.viii.1976, P. Rude, on Franseria (1♂, EMEC) , Fillmore , 3.ii.1970, R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle (1♂, UCR) , Ventura, R. D. Goeden & D.W. Ricker , insectary reared on Ambrosia psilostachys Decandolle , 16.iv.1969 (1♀, UCR) , 30.vi.1969 (1♀, UCR) , Port Hueneme , R. D. Goeden & D.W. Ricker, insectary reared on Ambrosia psilostachys Decandolle , 5.ii.1970 (1♀, UCR) , 15.iv.1969 (1♀, UCR) , Yolo Co., Woodland , 28.viii.1953, A. T. McClay (2♀, UCD) , Davis , 4.viii.1955, E.I. Schlinger (1♂, UCD) , Davis , 23.iv.1966, R.O. Schuster (1♂, UCD) , Montana: Mineral Co., St. Regis , 770m, 28.viii.1981, P.H. Arnaud, Jr. (1♂, CASC) , Columbia , Malaise trap, 7 AM –7 AM, 18.v.1970, F.D. Parker (1♂, UCD) .

Additional material examined, “brassicae 2” [on Foeniculum ]: USA. California: Solano Co., Cordelia , 24.vii.1959, P.H. Arnaud, Jr., coll. at flowers Foeniculum vulgare (L.) Gaertn. (1♂, USNM) .

Additional material examined, “brassicae 3”: USA. California:. San Bernardino Co., Crestline , 4.vi.1947, A.L. Melander (1♂, USNM) .

Additional material examined, Lantana -reared material: USA. California: HuntngtnBon [?], “”4/6/45”, A.L. Melander (1♂, USNM) , San Luis Obispo Co., San Luis Obispo City, on Lantana sp. , 20.iii.1997 (6♂ 4♀, CSCA) , Riverside, Riverside, A.L. Melander, 25.v.1942 (1♂, USNM) , Riverside, CR Picnic Hill, 30.iv.1997, M. Gates, ex. surface blotch mine on Lantana camara (1♂, USNM) , Riverside, UCR campus, M. Gates, ex. upper surface blotch mine on Lantana camara , 29.iv.1998 (5♀, USNM) , 8.v.1998 (1♂ 1♀, USNM) .

Comments. The material here recovered from Lantana is the second instance of Liriomyza brassicae reared from the family Verbenaceae , with Palacios et al. (2008) also finding it on Lantana camera L. in Mexico. The adult collected on Foeniculum is the first instance of this species on Umbelliferae, although rearing should confirm any potential host association. Morphologically, the Lantana -reared specimens are most obviously characterized by a slightly larger body size, a very narrow yellow line along the posterior margin of the scutum, a distiphallus that is more strongly constricted medially and wider distally (base only half width of distal section) and vein dm-cu is sometimes incomplete. The Foeniculum -collected male is darker overall, there are two closely spaced spines on the surstylus, and the distiphallus is dorsoventrally compressed basally and nearly parallel-sided in ventral view. Although L. brassicae is known to be quite polyphagous, these unusual plant associations (at least the confirmed Lantana records) in combination with the variant morphology suggest that this species may represent a species complex and/or collection of host races. This possibility is consistent with results from Tavormina (1982) who documented host-associated behavioural differences and strong host-associated selection in L. brassicae feeding on three sympatric host plant species.

The only other Agromyzidae known from Foeniculum are Melanagromyza foeniculi Spencer , Phytomyza ferulae Stackelberg and Ptochomyza czernyi (Strobl) , and the only Agromyzidae known from Lantana are Calycomyza bahamarum Spencer , C. lantanae Frick , L. trifolii , Ophiomyia camarae Spencer , O. lantanae (Froggatt) and O. legitima Spencer ( Perkins & Swezey 1924, Spencer 1990, Benavent-Corai et al. 2005).

In addition to this material, there is one male (“ brassicae 3”) that also has two subapical spines on the surstylus and a distiphallus that is nearly parallel-sided in ventral view. It is more distinct externally, however, in that the face is brown and the body is slightly darker. The host is unknown. Although there is potentially enough evidence to provide separate specific status to these three atypical groups, it seems preferable to gather additional biological data and material appropriate for molecular analysis before any taxonomic decisions are made, particularly considering the agricultural significance of this species.

In the United States, a similar phallus is found in Liriomyza baccharidis and L. conclavis , both of which have a predominantly dark pleuron, but the former (also found on Asteraceae ) has a dark face and a C-shaped surstylus. The latter has a relatively large first flagellomere, two long, subapical spines on the surstylus and a large, dorsallydirected apical chamber on the distiphallus. In Canada, a similar phallus is found in the Asclepias (Asclepiadaceae) feeders L. asclepiadis Spencer and L. peleensis Spencer , but the former is slightly larger (wing length 1.8–2.2mm), the distiphallus is broader past a strong medial constriction, and the apical chamber is truncated apically and dorsally and is more strongly angled. The latter is much darker with a narrower distiphallus.

Figures

FIGURES 54–60. Liriomyza brassicae (Riley), genitalia of “typical” male; 54: ejaculatory apodeme; 55: external components, anterior; 56: external components, posterior; 57: external components, ventral; 58: external components, left lateral; 59: phallus, left lateral; 60: phallus, ventral.

FIGURES 61–68. Figs 61–63: Liriomyza brassicae (Riley), genitalia of “brassicae 2” male collected on Foeniculum; 61: external components, ventral; 62: phallus, ventral; 63: phallus, left lateral. Figs 64–66: L. brassicae, genitalia of “brassicae 3” male; 64: external components, ventral; 65: phallus, ventral; 66: phallus, left lateral. Figs 67–68: L. brassicae, genitalia of male reared from Lantana; 67: phallus, ventral; 68: phallus, left lateral.