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In this study, we describe Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov., leafminers of herbaceous wetland plants. The M. erythranthei larva is a true miner entering fresh leaves and excavating the tunnels, and the L. viribus larva inhabits vacated mines of M. erythranthei. M. erythranthei is widespread in North America, with collections from the Pacific coast to Pennsylvania, and L. viribus has been collected from Iowa and Oregon. We also describe the larva of a possible new species associated with these plants, which we refer to as Metriocnemus sp. “Oregon”. A key to the known larval stages of North American Metriocnemus is also provided. Along with providing a detailed account of the mining ecology of these new species, we discuss additional observations of mostly Orthocladiinae midges associated with aquatic and terrestrial plants. These include documenting the rearing of Metriocnemus eurynotus (Holmgren, 1883) from larvae feeding on Impatiens (Balsaminaceae) cotyledons, initially as leafminers and later externally. Larvae of M. eurynotus also were found feeding within mines of M. erythranthei on Veronica (Plantaginaceae) and were collected along with M. erythranthei larvae on leaves of Petasites (Asteraceae).

Psectrocladius (s.s.) vernalis (Malloch, 1915) Material examined. USA: IOWA: Winneshiek Co., Decorah, Van Peenan Spring at Van Peenan Park, 43.312834, -91.776010, 17.v.2022, em. 22.v.2022, J. van der Linden, ex Veronica sp. (1Ô, USNM). Biological notes. The rearing container in which this specimen emerged included a single mined Veronica leaf inhabited by multiple Metriocnemus erythranthei larvae and one or two M. eurynotus larvae. The photographed M. eurynotus larva (Fig. 12a) exited the mine a short time later and began wandering on the leaf surface; it was later preserved. A second unmined Veronica leaf containing no larvae was added to the container at one point, and the M. erythranthei larvae began mining it after exiting their original leaf. The Psectrocladius vernalis male emerged five days after the original mined leaf was collected, along with a male of M. erythranthei. It would seem that it was one of the larvae feeding together in the mine and was not recognized as distinct from the M. erythranthei larvae. Larvae of P. vernalis have been found to live on the surfaces of submerged plants, including Eurasian milfoil (Haloragaceae: Myriophyllum spicatum L.) and native water stargrass (Pontederiaceae: Heteranthera dubia (Jacq.) MacMill.), but their habits were not observed (Balci & Kennedy 2003)., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on page 62, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Balci, P. & Kennedy, J. H. (2003) Comparison of chironomids and other macroinvertebrates associated with Myriophyllum spicatum and Heteranthera dubia. Journal of Freshwater Ecology, 18, 235 - 247. https: // doi. org / 10.1080 / 02705060.2003.9664490"]}

Metriocnemus sp. “ Oregon ” (Figs. 7a–f) Material examined. USA: OREGON: Lane Co., Blue River, 44.1535, -122.328, 3.vi.2022, leg. M. W. Palmer, ex Claytonia sibirica (4 larvae, USNM; 5 larvae, ANC); same but ex Myosotis scorpioides (1 larva, USNM); same but ex Petasites frigidus (2 larvae, USNM; 11 larvae, ANC). Larva (n = 7). Total length 4.5–6.0, 5.3 mm. Head 288–364, 339 μm long, 429–579, 458 μm wide. Coloration (Fig. 5a). Head capsule dark brown. Occipital margin much darker in contrast to the remainder of the head. Abdomen greenish yellow with a bluish pattern on the 1 st and 2 nd segments. Head. Antenna short, 5 segmented; antennal segments in μm: 43, 11–14, 3–4, 4–5, 5; 1 st antennal segment L/ W 1.2; ring organ located at mid-section of 1 st segment; AR 1.7–1.8, blade subequal to flagellum; blade 23–25, 24 μm long (Fig. 5b). SI trifid with lateral branches shorter, SII-SIII simple (Fig. 5c). Labral lamella comb-like (Fig. 5c). Premandible dark, with 2 basal and 2 inner teeth, 72–88, 74 μm long; brush well-developed (Fig. 5c). Mandible dark, apical tooth shorter than combined width of 4 inner teeth; seta subdentalis narrow reaching the base of basal inner teeth; setae interna with 7 branches, the apex of branches furcate (Fig. 5d), mandible 151–170, 161 μm long. Mentum dark, with wide bifid median tooth and 5 pairs of lateral teeth, median teeth sit much lower and are much smaller than 1 st lateral teeth; ventromental plate long running parallel to the lateral edge of mentum, reaching well beyond the base of last lateral tooth; seta submenti just posteriad to mentum aligned with 3 rd lateral tooth (Fig. 5e); mentum 94–116, 106 μm long and 110–126, 117 μm wide, ventromental plate 68–77, 74 μm long. Postmentum 132–157, 147 μm long. Abdomen. Posterior parapods wider than long, bearing around 15 simple dark claws, posterior parapod 157– 221, 174 μm long and 1166–239, 200 μm wide (Fig. 5f). Procercus almost as long as wide, bearing 5 apical setae, procercus 17–27, 21 μm long and 18–29, 24 μm wide, apical setae 107–118, 110 μm long. Four wide and conical anal tubules are present, anal tubules 177–234, 205 μm long. Diagnostic characters. The larva of Metriocnemus sp. “ Oregon ” can be separated from other related species by the combination of the following characteristics: Antenna short, 1 st antennal segment L/ W 1.2, ring organ located at mid-section of 1 st antennal segment, AR 1.7–1.8, blade subequal to flagellum; SI trifid with mid-branch longer; premandible with a well-developed brush; mentum dark, with wide bifid median tooth sitting much lower and are much smaller than 1 st lateral teeth, ventromental plate reaching well beyond the base of last lateral tooth, seta submenti aligned with 3 rd lateral tooth; posterior parapods wider than long, anal tubules semicircular. Biological notes. Larvae were collected along with leaf mines and larvae of Metriocnemus erythranthei on Claytonia sibirica, Myosotis scorpioides, and Petasites frigidus. The larvae in these samples were not observed closely, but on the Petasites, which was in the spray zone of a dripping rock seep, MWP noted that the dominant species had a dark band in the thoracic segments of the body, evidently referring to Metriocnemus sp. “ Oregon.” On this host, the mines were rather short, and the only larvae seen were window-feeding on the upper leaf surface (i.e., feeding externally and leaving the lower epidermis intact). Only leaves with mines were collected on the other two hosts, but these leaves also included larvae wandering externally. Thus, the degree to which M. sp. “ Oregon ” is a leafminer requires further investigation, and if it does feed in mines, it may be that it only does so within those initiated by M. erythranthei. Along with M. erythranthei and M. eurynotus, the Petasites collection also included two other chironomids: one larva of Boreochlus persimilis (Johannsen, 1926) (Podonominae) and one pupa of Orthocladius (Eudactylocladius) dubitatus Johannsen, 1942 (the adult female apparently emerged after the pupa was placed in ethanol). Boreochlus larvae have been previously reported to inhabit mosses in springs and small streams (Epler 2001), and Sublette et al. (1998) stated that those of O. dubitatus are probably madicolous (inhabiting thin films or sheets of water in streams, on vertical rock faces, etc.)., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on page 54, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Epler, J. H. (2001) Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. A guide to the taxonomy of the midges of the southeastern United States including Florida. North Carolina Department of Environment and Natural Resources Division of Water Quality, St. Johns River Water Management District, Palatka, Florida, iv + 530 pp.","Sublette, J. E., Stevens, L. E. & Shannon, J. P. (1998) Chironomidae (Diptera) of the Colorado River, Grand Canyon, Arizona, USA, I: systematics and ecology. The Great Basin Naturalist, 58, 97 - 146. https: // doi. org / 10.5962 / bhl. part. 12137"]}

Chironomus sp. Material examined. USA: OREGON: Lane Co., Blue River, 44.1541, -122.324, 5.iv.2022, M.W. Palmer, extracted from leaves of Sparganium emersum (3 larvae, ANC). Biological note. The larvae were found mining in submerged portions of leaves of Sparganium emersum Rehmann (Typhaceae) in a beaver pond. The mines were only externally visible in transmitted light., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on page 64, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232

A key to the known fourth instar larval stages of North American Metriocnemus van der Wulp 1. Mentum with 4 small median teeth and 5 pairs of lateral teeth (Epler 2001: page 7.86, couplet 2(1’))................................................................................................... M. knabi Coquillett, 1904 - Mentum with bifid or simple median tooth and 4 to 5 pairs of lateral teeth........................................ 2 2. Mentum with simple median tooth and 5 pairs of lateral teeth (Saether 1989: Figure 12G)...... M. ursinus (Holmgren, 1869) - Mentum with bifid median tooth and 4 to 5 pairs of lateral teeth................................................ 3 3. Mentum with a wide bifid median tooth and 4 pairs of lateral teeth (Fig. 3h)...................... M. erthranthei sp. nov. - Mentum with narrow bifid median tooth and 5 pairs of lateral teeth.............................................. 4 4. Median tooth of the mentum is comparatively small, sitting much lower than lateral teeth............................ 5 - Median tooth of the mentum comparatively well-developed, sitting slightly lower or higher than lateral teeth............ 6 5. Basal antennal segment no longer than wide (Saether 1989: Figure 12B). Labral lamella and SI simple (Saether 1989: Fig. 12D). Median tooth of mentum sunken (Epler 2001: page 7.86, couplet 3(2’); Orendt & Bendt 2021: page 119, couplet D7; Saether 1989: Fig. 12H). Anal tubules long and slender (Saether 1989: Fig. 12J)..................... M. fuscipes (Meigen, 1818) - Basal antennal segment longer than wide (Fig. 7b). Labral lamella comb-like and SI trifid (Fig. 5c). Median tooth of mentum not sunken (Fig. 7e). Anal tubules short and wide (Fig. 7f)......................................... M. sp. “ Oregon ” 6. Antenna 4 segmented (Cranston & Judd 1987: Fig. 4a). Basal antennal segment L/W> 4. Median tooth of mentum stands higher than 1 st lateral teeth. Setae submenti aligned with the 3 rd lateral teeth of mentum (Cranston & Judd 1987: Fig. 4c)............................................................................ M. yaquina Cranston & Judd, 1987 - Not with the above combination of characters............................................................... 7 7. Abdominal segments with long setae, each half as long as the segment bearing it.................... M. sp. A Epler, 2001 - Abdominal segments without long setae................................................................... 8 8. Procercus as wide as long or wider than long.AR 1.7–1.8. Basal antennal segment L/W: 2.3 (Th. Bendt & H.K.M. Moller Pillot, in litt.)................................................................................ M. tristellus agg. * - Not with the above combination of characters............................................................... 9 9. Median bifid tooth of mentum as high as the 1 st lateral teeth (Saether, 1989: Fig. 6E). Basal antennal segment L/ W 2.5 –2.6. Head length 0.44–0.45 mm................................................................. M. brusti Saether, 1989 - Median bifid tooth of mentum clearly lower than 1 st lateral teeth. Basal antennal segment L/ W 1.1 –2.4. Head length 0.13–0.47 mm............................................................................................... 10 10. Basal antennal segment L/ W 1.1. AR 0.67. Mandible with 3 inner teeth (Epler 2001: page 7.162). Procercus with 4 anal setae.............................................................. M. sp. (= Orthocladiinae genus E, Epler, 2001) ** - Not with the above combination of characters.............................................................. 11 11. AR 1.5–1.8. Head length 0.28-0.31 mm (Saether, 1989)............................... M. albolineatus (Meigen, 1818) - AR 2.0–2.2. Head length 0.39–0.47 (Saether, 1989)................................ M. eurynotus (Holmgren, 1883) ***, Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 56-57, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Epler, J. H. (2001) Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. A guide to the taxonomy of the midges of the southeastern United States including Florida. North Carolina Department of Environment and Natural Resources Division of Water Quality, St. Johns River Water Management District, Palatka, Florida, iv + 530 pp.","Saether, O. A. (1989) Metriocnemus van der Wulp: a new species and a revision of species described by Meigen, Zetterstedt, Staeger, Holmgren, Lundstr ˆ m and Strenzke (Diptera: Chironomidae). Insect Systematics & Evolution, 19, 393 - 430. https: // doi. org / 10.1163 / 187631289 X 00528","Orendt, C. & Bendt, Th. (2021) Orthocladiinae sensu lato (Orthocladiinae, Prodiamesinae, Diamesinae, Podonominae, Buchonomyiinae, Telmatogetoninae) (Diptera: Chironomidae). Keys to Central European larvae with respect to macroscopic characters. DGL-Arbeitshilfe (DGL Tools) 1 - 2021. German Limnological Society (DGL) e. V., Essen, 140 pp.","Cranston, P. S. & Judd, D. D. (1987) Metriocnemus (Diptera: Chironomidae): an ecological survey and description of a new species. Journal of the New York Entomological Society, 95, 534 - 546."]}

Limnophyes viribus Namayandeh, Eiseman, Palmer & van der Linden sp. nov. (Figs. 9–11) LSID: urn:lsid:zoobank.org:act:53ef97e1-5c99-4e60-9847-d3e1e61aae0c Holotype. USA: OREGON: Lane Co., Blue River, 44.1535, -122.328, 5.iv.2022, em. 10–14.iv.2022, leg. M. W. Palmer, ex Erythranthe guttata complex (1Ô, USNM). Paratypes. USA: IOWA: Winneshiek Co., Decorah, Van Peenan Spring, Van Peenan Park, 43.312834, - 91.776010, 17.v.2022, em. 21.v.2022, leg. J. van der Linden, ex. Impatiens sp. (1Ô, USNM); OREGON: same data as holotype (5ÔÔ, 4♀♀, 1 larva, USNM). Other material examined. USA: OREGON: same data as holotype (9ÔÔ, 2♀♀, ANC). Etymology. The new species is named after a phoenix sculpture named “ Viribus,” which represents resiliency and rebirth, made by sculptor Jud Turner. The monument stands in the McKenzie River Corridor town of Blue River, the type locality, which was ravaged by the Holiday Farm Fire of 2020. The word is Latin and means “strength.” Description. Male (n = 9). Total length 1.8–2.0, 1.9 mm. Wing 1.3–1.4 mm long and 0.3–0.4 mm wide. Coloration. Head, thorax, legs, tergites, sternites, and hypopygium black. Wings and halters grey. Head. Antenna with 13 flagellomeres, last flagellomere with 6 sensilla chaetica, groove starts at third segment, AR 0.3–0.6, 0.5. Eyes bare, without dorsomedial extension. Temporal setae few, 2 outer verticals and 1 frontal. Tentorium 132–135, 133 μm long (Fig. 9a). Clypeus rectangular, 60 μm long and 114 μm wide, bearing 15 setae, setae 57–70, 63 μm long. Palpal segment lengths (in μm): 27–29, 28; 32–39, 36; 65–68, 66; 49–57, 53; 70–108, 89. Third palpomere with 1 sensilla clavata. Thorax (Fig. 9b). Acrostichals 4; dorsocentrals 20–25, around 23 in a single row and remainder in post humeral region double rows; prealars 6–7; scutellars 6–7 in single row; 3–6 lanceolate humerals; 6–8 lanceolate prescutellars; 10–14, 12 antepronotals; 5 posterior anepisternals II; 3 median anepisternals II; 2 epimeron II; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical. Wing (Fig. 9c). Brachiolum with 1 seta. Squama bare. R with 11 setae, R 1 with 4 setae, other veins bare. Costa extension 45 μm. Anal lobe not projecting. Microtrichia visible at 10 ×. Legs. Fore tibia spur 37–45, 41 μm long, mid tibia spurs 20–24, 22 and 19 μm long, hind tibia spurs 42–50, 46 and 14–18, 16 μm long, hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 3. Hypopygium (Fig. 9d). Tergite IX with around 4 setae close to the base of anal point. Anal point extremely short, almost receded, wide with apex rounded; anal point 7–12, 10 μm long and 17–26, 21 μm wide. Virga consists of single long mid-spine with around 4 shorter lateral spines, the main spine 25–32, 28 μm long 7. Sternapodeme transverse with well-developed oral projections; sternapodeme 74–91, 81 μm long. Phallapodeme 33–42, 38 μm long. Inferior volsella a large triangular lobe with narrow apex; covered in numerous simple setae. Gonostylus large, expanded apically and with small spine-like distal outer projection, 60–77, 70 μm long; crista dorsalis large, overarching the apex of gonostylus. Gonocoxite 110–123, 116 μm long. HR 1.5–1.8, 1.7, HV 2.5–2.6. Female (n = 2). Total length 1.7–1.9, 1.8 mm. Wing 1.2 mm long and 0.37–0.43, 0.40 mm wide. Coloration. Same as the male. Head (Fig. 10a). Antenna with 5 flagellomeres, last flagellomere with 8 sensilla chaetica, 1 st –4 th segments each with 2 sensilla chaetica, AR 0.4–0.7, 0.5. Eyes bare. Temporal setae 3–4 including 2–3 outer vertical and 1 frontal. Tentorium 120–123, 122 μm long. Clypeus rectangular, 65–82, 74 μm long and 96–110, 104 μm wide, bearing 20 setae, setae 53–63, 59 μm long. Palpal segment lengths (in μm): 27–31, 29; 31–34, 32; 57–71, 64; 61; 72–90, 81. Thorax (Fig. 10b). Acrostichals 4; dorsocentrals 26, around 23 in a single row and remainder in post humeral region double rows; prealars 5; scutellars 7 in single row; 8 lanceolate humerals; 8 lanceolate prescutellars; 6 antepronotals; 4 posterior anepisternals II; 3 epimeron II; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical. Wing (Fig. 10c). Brachiolum with 1 seta. Squama bare. R with 9–13, 11 setae; R 1 with 5–7 setae; R 4+5 12–15 setae; other veins without setae. Costa extension 27–33 μm. Microtrichia visible at 10 ×. Legs. Hind and mid femur with keel. Fore tibia spur 24 μm long, mid tibia spurs 18–20, 19 and 13–17, 15 μm long, hind tibia spurs 40–47, 45 and 16–17 μm long; hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 4. Genitalia (Figs. 10d–e). Seminal capsules comparatively large, 66–70, 68 μm long, and 49–57, 53 μm wide, semi-circular, spermathecal ducts with loops, with well-developed bulb (Fig. 10d). Notum 154–174, 158 µm long. Gonapophysis VIII divided into ventrolateral and thin dorsomesal lobe (Fig. 10d).Apodeme lobe distinct. Gonocoxite developed with around 8–9 setae (Fig. 10e). Tergite IX undivided. Cercus small, crescent-shaped, 60–61 µm long, and 31–41, 36 µm wide (Fig. 10e). Pupa. At present not known. Pupal exuviae could not be retrieved from the leaves. Larva (n = 1). Total length 3.0 mm. Head 231 μm long and 267 μm wide. Coloration (Fig. 11a). Head capsule yellow with postmentum region darker. Occipital margin much darker in contrast to the remainder of the head. Abdomen bluish grey with patches of white. Head. Antenna 5 segmented; segments length in μm: 30, 11, 2, 5, 3; ring organ closer to the apex of the basal segment; blade as long as the flagellum, blade 21 μm long (Fig. 11b); AR 1.4. Labral SI serrate, SII–SIII simple (Fig. 11c). Premandible wide and bifid, dark apically, 49 μm long (Fig. 11c). Mandible dark, apical tooth shorter than combined width of 3 inner teeth; seta subdentalis very small; setae interna with several very long branches (Fig. 11d), mandible 86 μm long. Mentum dark, with large bifid median tooth and 5 pairs of lateral teeth, median tooth 2.8 × the 1 st lateral teeth; seta submenti posteriad to mentum aligned with 2 nd lateral tooth (Fig. 11e); mentum 57 μm long and 68 μm wide; ventromental plate 39 μm long, and 14 μm wide, large, slightly reaching beyond the margin of mentum. Abdomen. Posterior parapods 65 μm long and 40 μm long, bearing around 12 simple dark claws (Fig. 11f). Procercus 23 μm long and 17 μm wide, bearing 6 apical setae, apical setae 217 μm long; supraanal setae 144 μm long; four anal tubules with constriction, anal tubules 91 μm long (Fig. 11f). Diagnostic characters. Limnophyes viribus can be separated from other related species by the combination of the following characteristics: The adult male is characterized by AR of 0.3–0.6; 3–6 lanceolate humerals; 6– 8 lanceolate prescutellars; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical; anal point extremely short, almost receded; virga consists of single long mid-spine with around 4 shorter lateral spines; inferior volsella a large triangular lobe with narrow apex; gonostylus large, expanded apically with small spine-like distal outer projection. The adult female is characterized by AR of 0.4–0.7; 8 lanceolate humerals; 8 lanceolate prescutellars; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical; seminal capsules comparatively large. The larva is characterized by AR 1.4; premandible wide and bifid; setae interna of the mandible with several very long branches; mentum with large bifid median tooth and 5 pairs of lateral teeth; procercus bearing 6 apical setae; supraanal setae long; anal tubules longer than posterior parapods. Taxonomic remarks. The adults of L. viribus resemble Limnophyes pilicistulus Saether, 1975. The two species are probably related and form a sister group. Adults of L. viribus can be separated from L. pilicistulus by the lower number of temporal setae, higher number of lanceolate humerals and prescutellars. Additionally, the adult male of L. viribus has a virga with more branches, and a gonostylus with more overarching crista dorsalis, and a small spinelike distal outer projection. The adult female of L. viribus has longer cercus and notum and shorter seminal capsules compared to L. pilicistulus. Biological notes. The larvae are apparently secondary inhabitants in leaf mines of Metriocnemus species, as with the undetermined Limnophyes species discussed below. The single Iowa specimen emerged in a batch rearing of M. eurynotus larvae feeding on Impatiens cotyledons. Four days earlier, in another rearing container, a mine with a larva of M. eurynotus had been found to also contain a much smaller larva that appeared uniformly dark and may have been an individual of L. viribus (photos at https://bugguide.net/node/view/2126834). Also in Iowa, a female very likely representing L. viribus was reared from a collection of M. erythranthei larvae mining Veronica leaves (see below under “ Limnophyes spp. ”). The Oregon specimens were reared along with M. erythranthei from plants of the Erythranthe guttata complex, although it was not entirely clear whether they emerged from leaves or the muck surrounding the roots., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 57-61, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232

Metriocnemus eurynotus (Holmgren, 1883) (Figs. 6–7) Material examined. USA: IOWA: Winneshiek Co., Decorah, Van Peenan Spring at Van Peenan Park, 43.312834, -91.776010, 14.iv.2022, em. 6.v.2022, leg. J. van der Linden, ex thallose liverwort (1Ô, 1 pupa, 1 larva, ANC); same location, 10.v.2022, preserved 16.v.2022, leg. J. van der Linden, ex Impatiens sp. (1 larva, USNM); same but preserved 21.v.2022 (2 pupae, ANC); same but em. 21.v.2022, (1♀, ANC); same but em. 23.v.2022 (2♀, 2 pupae, 2 larvae, ANC); same but em. by 27.v.2022 (2♀♀, ANC); same location, 17.v.2022, em. 21.v.2022, leg. J. van der Linden, ex Impatiens sp. (1Ô, 1 pupa, 1 larva, USNM); same but em. 23.v.2022 (1♀, ANC); same but em. by 27.v.2022 (2♀♀, 1 pupa, ANC); same location, 17.v.2022, leg. J. van der Linden, ex Veronica sp. (1 larva, USNM); same but em. 29.v.2022 (1♀, 2 pupae, 1 larva, USNM); OREGON: Lane Co., Blue River, 44.1535, -122.328, 3.vi.2022, leg. M. W. Palmer, ex Petasites frigidus (2 larvae, USNM). Biological notes. It appears that nothing has been published previously about the larval habits of this Holarctic species (Saether 1989, 1995). John van der Linden (JvdL) was examining 5- to 10-cm tall sprouts of jewelweed (Balsaminaceae: Impatiens sp.; very likely I. capensis Meerb.) on the margin of a rocky spring-fed pool in Iowa in early May when he noticed mines in cotyledons of some of the plants (Fig. 6a). Chironomid larvae were present inside the mines and on the surfaces of the cotyledons and appeared to move freely between these niches (Figs. 6b–h). The mines, which were only observed in Impatiens cotyledons and not in the developing true leaves, consisted of short, irregular tunnels into the tissue emanating from a central blotch. Tears or holes in one or both epidermises in the central blotch area allowed larvae to enter and exit the mines. In some older mines, the central area’s epidermis had fallen away completely, resulting in a hole in the cotyledon. In captivity, larvae were observed to move around in the mines and feed on mesophyll. The mines contained sparsely scattered green or greenish-brown pellets or rods of frass. One feeding larva with similarly colored material in its gut was observed depositing excrement in the mine. As many as five larvae were observed inside or on the edges of one mined area on a single cotyledon (video at https:// youtu.be/Tt4 SYMH 3U5U). Larvae interacted vigorously, especially upon physical contact or very close proximity, to which they responded with thrashing movements or by appearing to bite or nip each other’s bodies (Fig. 6i). Most captive larvae soon exited the mines permanently. They switched to feeding externally on the cotyledons (Fig. 6j). In containers holding multiple larvae, two or three individuals fed communally on a cotyledon and, in some cases, consumed it entirely before reaching maturity. Full-grown larvae pupated exposed on the cotyledon remnants or on the moist paper towel bedding or sides of the containers (Fig. 6k). Nine adults emerged (Fig. 6l). Several other individuals reached pupation. However, they failed to emerge as adults, with some darkened, mature pupae crawling around the rearing container actively, only to perish a short time later. Larvae of M. eurynotus were not observed to initiate new mines in pristine cotyledons. Although no other insects were observed feeding on or in them, we cannot exclude the possibility that there was some initial damage to the cotyledons that allowed the larvae to enter them. A week after the larvae were first discovered, more M. eurynotus larvae were found on leaves of Veronica sp. in the same rocky spring, both moving around on the leaf surfaces and feeding within leaf mines of M. erythranthei (Fig. 12a; videos at https://youtu.be/n9J6RJ0-DnI and https://youtu.be/qQ6k2eWBz_I). One male was also reared from a thallose liverwort collected from this site in midApril, although the larva was never actually observed in this case. There were mines in the liverwort, at least some of which appeared to be agromyzid in origin. It is conceivable that the M. eurynotus individual fed as a larva inside these. Before starting the rearing, the undersides and rhizoids of the thalli were thoroughly washed and massaged in water to remove the substrate. The pupa of this individual was found loose in the rearing container on 5 May, and examination of the thalli at this time revealed that a few of them showed evidence of heavy external feeding; no possible source for this was found other than the M. eurynotus, and the damage was similar in appearance to the external feeding on the jewelweed cotyledons. In Oregon, Mike W. Palmer (MWP) collected larvae of M. eurynotus on leaves of Petasites frigidus along with M. erythranthei and the Metriocnemus species discussed below., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 51-54, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Saether, O. A. (1989) Metriocnemus van der Wulp: a new species and a revision of species described by Meigen, Zetterstedt, Staeger, Holmgren, Lundstr ˆ m and Strenzke (Diptera: Chironomidae). Insect Systematics & Evolution, 19, 393 - 430. https: // doi. org / 10.1163 / 187631289 X 00528"]}

Limnophyes viribus Namayandeh, Eiseman, Palmer & van der Linden sp. nov. (Figs. 9–11) LSID: urn:lsid:zoobank.org:act:53ef97e1-5c99-4e60-9847-d3e1e61aae0c Holotype. USA: OREGON: Lane Co., Blue River, 44.1535, -122.328, 5.iv.2022, em. 10–14.iv.2022, leg. M. W. Palmer, ex Erythranthe guttata complex (1Ô, USNM). Paratypes. USA: IOWA: Winneshiek Co., Decorah, Van Peenan Spring, Van Peenan Park, 43.312834, - 91.776010, 17.v.2022, em. 21.v.2022, leg. J. van der Linden, ex. Impatiens sp. (1Ô, USNM); OREGON: same data as holotype (5ÔÔ, 4♀♀, 1 larva, USNM). Other material examined. USA: OREGON: same data as holotype (9ÔÔ, 2♀♀, ANC). Etymology. The new species is named after a phoenix sculpture named “ Viribus,” which represents resiliency and rebirth, made by sculptor Jud Turner. The monument stands in the McKenzie River Corridor town of Blue River, the type locality, which was ravaged by the Holiday Farm Fire of 2020. The word is Latin and means “strength.” Description. Male (n = 9). Total length 1.8–2.0, 1.9 mm. Wing 1.3–1.4 mm long and 0.3–0.4 mm wide. Coloration. Head, thorax, legs, tergites, sternites, and hypopygium black. Wings and halters grey. Head. Antenna with 13 flagellomeres, last flagellomere with 6 sensilla chaetica, groove starts at third segment, AR 0.3–0.6, 0.5. Eyes bare, without dorsomedial extension. Temporal setae few, 2 outer verticals and 1 frontal. Tentorium 132–135, 133 μm long (Fig. 9a). Clypeus rectangular, 60 μm long and 114 μm wide, bearing 15 setae, setae 57–70, 63 μm long. Palpal segment lengths (in μm): 27–29, 28; 32–39, 36; 65–68, 66; 49–57, 53; 70–108, 89. Third palpomere with 1 sensilla clavata. Thorax (Fig. 9b). Acrostichals 4; dorsocentrals 20–25, around 23 in a single row and remainder in post humeral region double rows; prealars 6–7; scutellars 6–7 in single row; 3–6 lanceolate humerals; 6–8 lanceolate prescutellars; 10–14, 12 antepronotals; 5 posterior anepisternals II; 3 median anepisternals II; 2 epimeron II; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical. Wing (Fig. 9c). Brachiolum with 1 seta. Squama bare. R with 11 setae, R 1 with 4 setae, other veins bare. Costa extension 45 μm. Anal lobe not projecting. Microtrichia visible at 10 ×. Legs. Fore tibia spur 37–45, 41 μm long, mid tibia spurs 20–24, 22 and 19 μm long, hind tibia spurs 42–50, 46 and 14–18, 16 μm long, hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 3. Hypopygium (Fig. 9d). Tergite IX with around 4 setae close to the base of anal point. Anal point extremely short, almost receded, wide with apex rounded; anal point 7–12, 10 μm long and 17–26, 21 μm wide. Virga consists of single long mid-spine with around 4 shorter lateral spines, the main spine 25–32, 28 μm long 7. Sternapodeme transverse with well-developed oral projections; sternapodeme 74–91, 81 μm long. Phallapodeme 33–42, 38 μm long. Inferior volsella a large triangular lobe with narrow apex; covered in numerous simple setae. Gonostylus large, expanded apically and with small spine-like distal outer projection, 60–77, 70 μm long; crista dorsalis large, overarching the apex of gonostylus. Gonocoxite 110–123, 116 μm long. HR 1.5–1.8, 1.7, HV 2.5–2.6. Female (n = 2). Total length 1.7–1.9, 1.8 mm. Wing 1.2 mm long and 0.37–0.43, 0.40 mm wide. Coloration. Same as the male. Head (Fig. 10a). Antenna with 5 flagellomeres, last flagellomere with 8 sensilla chaetica, 1 st –4 th segments each with 2 sensilla chaetica, AR 0.4–0.7, 0.5. Eyes bare. Temporal setae 3–4 including 2–3 outer vertical and 1 frontal. Tentorium 120–123, 122 μm long. Clypeus rectangular, 65–82, 74 μm long and 96–110, 104 μm wide, bearing 20 setae, setae 53–63, 59 μm long. Palpal segment lengths (in μm): 27–31, 29; 31–34, 32; 57–71, 64; 61; 72–90, 81. Thorax (Fig. 10b). Acrostichals 4; dorsocentrals 26, around 23 in a single row and remainder in post humeral region double rows; prealars 5; scutellars 7 in single row; 8 lanceolate humerals; 8 lanceolate prescutellars; 6 antepronotals; 4 posterior anepisternals II; 3 epimeron II; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical. Wing (Fig. 10c). Brachiolum with 1 seta. Squama bare. R with 9–13, 11 setae; R 1 with 5–7 setae; R 4+5 12–15 setae; other veins without setae. Costa extension 27–33 μm. Microtrichia visible at 10 ×. Legs. Hind and mid femur with keel. Fore tibia spur 24 μm long, mid tibia spurs 18–20, 19 and 13–17, 15 μm long, hind tibia spurs 40–47, 45 and 16–17 μm long; hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 4. Genitalia (Figs. 10d–e). Seminal capsules comparatively large, 66–70, 68 μm long, and 49–57, 53 μm wide, semi-circular, spermathecal ducts with loops, with well-developed bulb (Fig. 10d). Notum 154–174, 158 µm long. Gonapophysis VIII divided into ventrolateral and thin dorsomesal lobe (Fig. 10d).Apodeme lobe distinct. Gonocoxite developed with around 8–9 setae (Fig. 10e). Tergite IX undivided. Cercus small, crescent-shaped, 60–61 µm long, and 31–41, 36 µm wide (Fig. 10e). Pupa. At present not known. Pupal exuviae could not be retrieved from the leaves. Larva (n = 1). Total length 3.0 mm. Head 231 μm long and 267 μm wide. Coloration (Fig. 11a). Head capsule yellow with postmentum region darker. Occipital margin much darker in contrast to the remainder of the head. Abdomen bluish grey with patches of white. Head. Antenna 5 segmented; segments length in μm: 30, 11, 2, 5, 3; ring organ closer to the apex of the basal segment; blade as long as the flagellum, blade 21 μm long (Fig. 11b); AR 1.4. Labral SI serrate, SII–SIII simple (Fig. 11c). Premandible wide and bifid, dark apically, 49 μm long (Fig. 11c). Mandible dark, apical tooth shorter than combined width of 3 inner teeth; seta subdentalis very small; setae interna with several very long branches (Fig. 11d), mandible 86 μm long. Mentum dark, with large bifid median tooth and 5 pairs of lateral teeth, median tooth 2.8 × the 1 st lateral teeth; seta submenti posteriad to mentum aligned with 2 nd lateral tooth (Fig. 11e); mentum 57 μm long and 68 μm wide; ventromental plate 39 μm long, and 14 μm wide, large, slightly reaching beyond the margin of mentum. Abdomen. Posterior parapods 65 μm long and 40 μm long, bearing around 12 simple dark claws (Fig. 11f). Procercus 23 μm long and 17 μm wide, bearing 6 apical setae, apical setae 217 μm long; supraanal setae 144 μm long; four anal tubules with constriction, anal tubules 91 μm long (Fig. 11f). Diagnostic characters. Limnophyes viribus can be separated from other related species by the combination of the following characteristics: The adult male is characterized by AR of 0.3–0.6; 3–6 lanceolate humerals; 6– 8 lanceolate prescutellars; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical; anal point extremely short, almost receded; virga consists of single long mid-spine with around 4 shorter lateral spines; inferior volsella a large triangular lobe with narrow apex; gonostylus large, expanded apically with small spine-like distal outer projection. The adult female is characterized by AR of 0.4–0.7; 8 lanceolate humerals; 8 lanceolate prescutellars; 8 preepisternals, 6 anteriorly clustered and diagonal, separated from 2 vertical; seminal capsules comparatively large. The larva is characterized by AR 1.4; premandible wide and bifid; setae interna of the mandible with several very long branches; mentum with large bifid median tooth and 5 pairs of lateral teeth; procercus bearing 6 apical setae; supraanal setae long; anal tubules longer than posterior parapods. Taxonomic remarks. The adults of L. viribus resemble Limnophyes pilicistulus Saether, 1975. The two species are probably related and form a sister group. Adults of L. viribus can be separated from L. pilicistulus by the lower number of temporal setae, higher number of lanceolate humerals and prescutellars. Additionally, the adult male of L. viribus has a virga with more branches, and a gonostylus with more overarching crista dorsalis, and a small spinelike distal outer projection. The adult female of L. viribus has longer cercus and notum and shorter seminal capsules compared to L. pilicistulus. Biological notes. The larvae are apparently secondary inhabitants in leaf mines of Metriocnemus species, as with the undetermined Limnophyes species discussed below. The single Iowa specimen emerged in a batch rearing of M. eurynotus larvae feeding on Impatiens cotyledons. Four days earlier, in another rearing container, a mine with a larva of M. eurynotus had been found to also contain a much smaller larva that appeared uniformly dark and may have been an individual of L. viribus (photos at https://bugguide.net/node/view/2126834). Also in Iowa, a female very likely representing L. viribus was reared from a collection of M. erythranthei larvae mining Veronica leaves (see below under “ Limnophyes spp. ”). The Oregon specimens were reared along with M. erythranthei from plants of the Erythranthe guttata complex, although it was not entirely clear whether they emerged from leaves or the muck surrounding the roots.

Metriocnemus erythranthei Namayandeh, Eiseman, van der Linden & Palmer sp. nov. (Figs. 1–5) LSID: urn:lsid:zoobank.org:act: 524CFBAA-C5C4-48B6-A219-A71614991FB8 Holotype. USA: OKLAHOMA: Woodward Co., Boiling Spring State Park; 2.v.2021, em. 10–11.v.2021, leg. E. LoPresti and K. Toll, ex Erythranthe glabrata (1Ô, USNM). Paratypes. Canada: BRITISH COLUMBIA: Cowichan Valley, 48.837266, -123.590338, 25.iii.2022, leg. F. McGhee, ex Erythranthe guttata (1 pupa, 1 larva, USNM); USA: CALIFORNIA: Sonoma Co., Bodega Bay, Campbell Cove; 31.v.2019, em. by 3.vi.2019, leg. K. Toll, ex Erythranthe guttata (1 larva, USNM); IOWA: Winneshiek Co., Decorah, Van Peenan Spring at Van Peenan Park, 43.312834, -91.776010, 17.v.2022, em. 22.v.2022, leg. J. van der Linden, ex Veronica sp. (1Ô, USNM); OKLAHOMA: same data as holotype (1Ô, 2♀, 2 pupae, 1 larva, USNM); OREGON: Lane Co., 44.235804, -122.85404, 9.v.2022, em. 14.v.2022, leg. J. Ward, ex Claytonia sibirica (1♀, 1 pupa, USNM); Lane Co., Blue River, 44.1535, -122.328, 3.vi.2022, leg. M. W. Palmer, ex Claytonia sibirica (1Ô, 1 pupa, 1 larva, USNM); same but ex Myosotis scorpioides (1 pupa, USNM); same but ex Petasites frigidus, adults em. 12–20.vi.2022 (3ÔÔ, 2♀♀, 1 pupa, 1 larva, USNM); PENNSYLVANIA: Berks Co., Blandon, 40.461258, -75.88125, 8.v.2022, preserved 14.v.2022, leg. C. Smith, ex Veronica anagallis-aquatica (1 larva, USNM). Other material examined. USA: CALIFORNIA: Santa Cruz Co., Bonny Doon, 31.x.2012; leg. C.S. Eiseman, ex Erythranthe moschata (1 larva, JHEC); IOWA: Winneshiek Co., Decorah, Twin Springs Park, 43.297623, - 91.815370, 4.iv.2022, em. 6.iv.2022, J. van der Linden, ex Veronica sp. (1Ô, 1 pupa, 1 larva, ANC); same but 6.iv.2022 (2 pupae, 3 larvae, ANC); same location, 17.v.2022, em. 22.v.2022, J. van der Linden, ex Veronica sp. (4ÔÔ, 4 pupae, 1 larva, ANC); OREGON: Lane Co., Blue River, 44.1535, -122.328, 6.iv.2022, leg. M. W. Palmer, ex Erythranthe guttata complex (6 pupae, 12 larvae, ANC); same collection, em. 10–14.iv.2022 (3ÔÔ, 1♀, ANC); same but 28.iv.2022 (16 larvae, ANC); same but 3.vi.2022, ex Stachys cooleyae (8 larvae, ANC); same but 3.vi.2022, adult em. ~ 8.vi.2022, ex Mentha × piperita ssp. citrata (1♀, 1 pupa, ANC); same but 3.vi.2022, adult em. ~ 8.vi.2022, ex Myosotis scorpioides (1Ô, 1 larva, ANC); same but 3.vi.2022, adults em. 12–20.vi.2022, ex Petasites frigidus (1Ô, 11♀♀, 6 pupae, 4 larvae, ANC); Blue River, 44.1507, -122.324, 28.iv.2022, leg. M. W. Palmer, ex Veronica americana (11 larvae, ANC); PENNSYLVANIA: Berks Co., Blandon, 40.461258, -75.88125, 8.v.2022, preserved 14.v.2022, leg. C. Smith, ex Veronica anagallis-aquatica (1 larva, ANC). Photographed leaf mines. Canada: BRITISH COLUMBIA: Capital Regional District, Langford, Goldstream Provincial Park, 48.483147, -123.551983, 13.viii.2022, L. Ragan, Claytonia sibirica (iNat 130692120); Cowichan Valley, 48.837266, -123.590338, 25.iii.2022, F. McGhee, Erythranthe guttata (iNat 110361786); USA: ALASKA: Sitka Co., Sitka, 57.08361, -135.282176, 15.vii.2021, J. Goff, Erythranthe guttata (iNat 87446898); Sitka Co., 57.123319, -135.313118, 4.viii.2022, M. Goff, Erythranthe guttata (iNat 129592195); CALIFORNIA: Marin Co., Golden Gate National Recreation Area, 37.830228, -122.505225, 12.vi.2022, C. Chang, Erythranthe Sect. Simiolus (iNat 121533464); Napa Co., McLaughlin Preserve, 38.856449, -122.402827, 21.vi.2016, E. LoPresti, Erythranthe guttata (iNat 69586931); Santa Clara Co., San Jose, 37.13310, -121.778717, 29.iv.2022, M. Vonshak, Veronica (iNat 115791054); Sonoma Co., Sebastopol, Morelli Ln, 38.430297, -122.95137, 8.ii.2022, K. Toll, Erythranthe guttata (iNat 106618091); OREGON: Lane Co., 44.153482, -122.329189, 27.iii.2021, M. Palmer, Erythranthe?nasuta (guttata complex) (iNat 109961926); 43.942162, -123.892629, 7.v.2022, J. Ward, Claytonia cf. sibirica (iNat 116020762); 43.637107, -122.617126, 13.vii.2022, J. Ward, Claytonia sibirica (iNat 126161890); Lincoln Co., Cape Perpetua Overlook, 44.287479, -124.110256, 25.vi.2022, J. Ward, Claytonia sibirica (larvae observed, but only empty mines in photos) (iNat 123640622); Linn Co., 44.378202, -122.000636, 22.ix.2022, J. Ward, Erythranthe moschata (iNat 136120938); Marion Co., 44.883958, -122.619387, 7.vii.2022, J. Ward, Claytonia sibirica (empty mines) (iNat 125263360); PENNSYLVANIA: Berks Co., Peters Creek Spring, 27.x.2021, C. Smith, Veronica (iNat 100832107, 100832147, 100832157, 100832185); WASHINGTON: King Co., Vashon, 47.390545, -122.489901, 24.iv.2020, H. Parker, Myosotis (iNat 44625412); same but 3.v.2020 (iNat 52704648, 59353928, 59355328); WISCONSIN: Waukesha Co., Eagle Spring and Fen, 42.924285, -88.468514, 24.ix.2014, D. Carter, Erythranthe geyeri (iNat 893113). Etymology. The new species is named after Erythranthe (monkeyflowers), one of the plant genera commonly inhabited by the larvae. Description. Male (n = 11). Total length 1.9–2.5, 2.2 mm. Wing 1.8–1.9 mm long and 0.45 mm wide. Coloration. Head, thorax, legs, tergites, sternites II–VIII, and hypopygium dark brown. Wings, halters, and sternite I greyish. Head. Antenna with 12 flagellomeres, last flagellomere with 8 sensilla chaetica, second to third segments each with 2 sensilla chaetica, groove starts at third segment, AR 0.9. Eyes bare, with wedge-shaped dorsomedial extension. Temporal setae 32 in several rows. Tentorium with bulging apex, 183–194, 189 μm long (Fig. 1a). Clypeus rectangular, 83–96, 89 μm long and 121–154, 138 μm wide, bearing 26 setae, setae 73–88, 81 μm long. Palpal segment lengths (in μm): 72–76, 74; 51; 129–159, 144; 80–95, 88; 141–167, 154. Third palpomere with 3 sensilla clavata. Thorax (Fig. 1b). Acrostichals 25–30, 27; dorsocentrals 79–109, 94 in multiple rows; prealars 21–24, 22; scutellars 42–44, 43 in three rows; supraalars 2–3. Antepronotal lobes developed, with a gap, 14–16, 15 lateral setae. Wing (Fig. 1c). Brachiolum with 10 setae. Squama with 18 setae. Sc with 30–33, 32; R with 32–42, 37 and R 1 with 35–43, 39 setae, R 4+5 with 50–54, 52 setae, and M with 25–28, 27 setae, other veins bare. Costa well-extended, extension 77 μm. R 4+5 ends just above M 3+4. Anal lobe not projecting (Fig. 1c). Legs. Tibia of all legs with long sparse beard; hind and mid femur with keel. Pulvilli very small. Fore tibia spur 62–72, 67 μm long, mid tibia spurs 34–39, 36 and 33–34 μm long, hind tibia spurs 55–65, 60 and 28 μm long, hind tibia comb with around 12 spines. 2 pseudospurs on ta 1 of mid and hind legs. Lengths and proportions of legs as in Table 1. Hypopygium (Fig. 1d). Tergite IX with around 14 long setae close to the base of anal point. Anal point short, narrowly triangular, apex rounded; anal point 32–46, 39 μm long and 17–23, 20 μm wide. Virga present, consisting of about 8–9 spines, 38–46, 42 μm long. Sternapodeme nearly straight, 124–136, 130 μm long. Phallapodeme 89–95, 92 μm long. Inferior volsella located anteriorly on gonocoxite, slightly bulging, covered in numerous simple setae. Gonostylus 113–121, 117 μm; crista dorsalis a preapical short triangle. Gonocoxite 231–246, 238 μm long. HR 2, HV 2.5–2.9, 2.7. Female (n = 5). Total length 2.8–3.1, 3.0 mm. Wing 1.8–1.9 mm long and 0.6 mm wide. Coloration. Humeral, anepisternal, preepisternum, and posnotum light brown, remainder same as the male. Head. Antenna with 5 flagellomeres, last flagellomere 60–73, 67 μm long; last flagellomere with 12 sensilla chaetica, 1 st –4 th segments each with 2 sensilla chaetica, AR 0.3 (Fig. 2a). Eyes bare, with short wedge-shaped dorsomedial extension. Temporal setae 43–50, 47 in several rows. Tentorium with bulging apex, 192–198, 195 μm long. Clypeus rectangular, 101–107, 104 μm long and 149–165, 157 μm wide, bearing 30–36, 33 setae, setae 79–100, 81 μm long. Palpal segment lengths (in μm): 61; 36–42, 39; 122–156, 139; 98–118, 108; 135–181, 158. Third palpomere with 1 sensilla clavata. Thorax. Acrostichals 34–40, 37; dorsocentrals 105–112, 109 in multiple rows; prealars 27–33, 30; scutellars 50– 54, 52 in two rows; supraalars 5.Antepronotal lobes developed, with a gap, 14–17, 15 lateral setae. Humeral pit small. Wing (Fig. 2b). Brachiolum with 9 setae. Squama with 19–22, 22 setae. R with 40–57, 50; R 1 with 27–40, 34 setae; R 4+5 with 50 setae, and M with 24–25 setae, other veins without setae. Costa well-extended, extension 45 μm. R 4+5 ends just above M 3+4. Anal lobe not projecting Legs. Hind-fore tibia with long sparse beard, hind and mid femur with keel. Pulvilli very small. Fore tibia spur 37–41, 39 μm long, mid tibia spurs 31–38, 34 and 30 μm long, hind tibia spurs 45–56, 51 and 25–33, 29 μm long, hind tibia comb with around 15 spines. 2 pseudospurs on ta 1 of mid and hind leg. Lengths and proportions of legs as in Table 2. Genitalia (Figs. 2c–d). Seminal capsules small, semi-circular, spermathecal ducts without loops, with small bulb (Fig. 2c), seminal capsule 67–68 µm long and 54–67, 61 µm wide. Notum 228–271, 249 µm long. Gonapophysis VIII divided into large ventrolateral and smaller dorsomesal lobe (Fig. 2c). Apodeme lobe distinct. Gonocoxite developed with around 24 setae (Fig. 2d). Tergite IX undivided (Fig. 2d). Cercus pediform, 152–156, 154 µm long, and 60 µm wide. Pupa (n = 8). Total length 4.5 mm. Coloration: Head and thorax brown. Abdomen golden. Cephalothorax. Frontal setae absent, frontal apotome rugose (Fig. 3a). Antennal sheet without pearls or spines above to pedicels. Thorax rugose, horn absent. Wing sheet nearly smooth, 1.1–1.2 mm long and 0.32–0.39, 0.35 mm wide. Abdomen (Figs. 3b–c). Tergite I with no posterior row of tubercles and no shagreens. Tergite II with no shagreens, a single row of posterior tubercles; tergites III–VIII with a single row of posterior tubercles; tergites III–VI with anterior crescent-shaped shagreens, becoming more prominent towards segment VI; tergites VII–VIII with shagreens more prominent anteriorly, becoming reduced from mid to posterior regions; tergite IX covered in shagreen (Fig. 3b). Sternites I, II, and VIII bare; sternites III–VII with posterior shagreens; sternite IX bare (Fig. 3c). Anal lobe 233–278, 256 µm long and 159–178, 171 µm wide; with two to three extremely short and hairlike macrosetae, 4.3–4.7, 4.5 µm long. Genital sac shorter than anal lobes, 163–167, 165 µm long and 105–122, 113 µm wide. Larva (n = 4). Total length 4.4 mm. Head 327–407, 379 μm long, 234–303, 268 μm wide. Coloration (Fig. 3d). Head capsule ventrally light brown, dorsally dark brown. Occipital margin much darker in contrast to the remainder of the head. Abdomen yellowish green. Head. Head L/ W 1.3 –1.5, 1.4. Antenna short, 5 segmented; 1 st antennal segment 1.5–2 × as long as wide; ring organ located at basal ¼ of 1 st segment; AR 1.3–1.5, 1.4, blade longer than the flagellum, blade 25–27, 26 μm long, B 2 6 μm long; antennal segments length in μm: 28–32, 8, 3.4–4, 3, 6–6.6 (Fig. 3e). SI divided into 4 branches, SIISIII simple (Fig. 3f). Premandible dark, with 2 basal and 2 inner teeth, 68–76, 72 μm long, brush developed (Fig. 3f). Mandible dark, apical tooth shorter than combined width of 4 inner teeth; seta subdentalis narrow reaching the base of basal inner teeth; setae interna with 7 branches, the apex of branches furcate (Fig. 3g), mandible 137–149, 141 μm long. Mentum dark, with wide bifid median tooth and 4 pairs of lateral teeth, ventromental plate running parallel to the lateral edge of mentum, reaching the base of last lateral tooth; seta submenti just posteriad to mentum aligned with 2 nd lateral teeth (Fig. 3h); mentum 86–98, 93 μm long and 105–142, 120 μm wide, ventromental plate 47–48 μm long and 10 μm wide. Abdomen. Posterior parapods well-reduced, much wider than long, bearing around 15 simple claws, posterior parapod 62 μm long and 140 μm wide. Procercus almost as long as wide, bearing 5 apical setae, procercus 19–20 μm long and 22–23 μm wide, apical setae 92–105, 98 μm long. Diagnostic characters. Metriocnemus erythranthei can be separated from other related species by the combination of the following characteristics: Adult male with AR 0.9; temporal setae 32 in several rows; tentorium with bulging apex; anal point short, moderately wide, and triangular, apex slightly rounded; virga consists of about 8–9 spines; inferior volsella located anteriorly and slightly bulging; crista dorsalis of gonostylus preapical, short and triangular. Adult female with AR 0.9; last flagellomere 67 μm long, last flagellomere/2 nd flagellomere 1.2; temporal setae 43–50 in several rows; seminal capsules small, semi-circular, spermathecal ducts without loops, with small bulb; gonapophysis VIII divided into large ventrolateral and dorsomesal lobe; apodeme lobe distinct; gonocoxite well-developed with around 24 setae; cercus pediform. Pupa with frontal setae absent; frontal apotome rugose; tergites I–II with no shagreens; tergites III–VIII with shagreens; tergites II–VIII with a single row of posterior tubercles. Sternites I, VII, and VIII bare; sternites III–VI with posterior shagreens; sternite IX covered in shagreen. Anal lobe with two to three extremely short and hairlike macrosetae; genital sac shorter than anal lobes. Larva with short antenna; basal antennal segment 1.5–2 × as long as wide; AR 1.4, blade longer than the flagellum; SI divided into 4 branches; premandible with 2 basal, 2 inner teeth and brush; mandible apical tooth shorter than combined width of 4 inner teeth; mentum with wide bifid median tooth and 4 pairs of lateral teeth; posterior parapods wellreduced. Biological notes (Figs. 4–5). Metriocnemus erythranthei mines leaves of at least three different monkeyflower species: Erythranthe glabrata (Kunth) G.L.Nesom in Oklahoma (Fig. 4a), E. guttata (DC.) G.L.Nesom (or E. guttata complex) in British Columbia, Oregon, and California (Fig. 4b), and E. moschata (Douglas ex Lindl.) G.L.Nesom in Oregon and California (Figs. 4c–d). Mines, probably representing this species, have been photographed in Alaska on E. guttata, as well as in Wisconsin on E. geyeri (Torr.) G.L.Nesom. In Oregon, Iowa, and Pennsylvania, M. erythranthei mines leaves of Veronica americana (Raf.) Schwein. ex Benth. and V. anagallis-aquatica L. (Plantaginaceae) (Figs. 4e–g, 5a–b), and in Oregon it has also been found to feed on Mentha aquatica L. and Stachys chamissonis var. cooleyae (A.Heller) G.A.Mulligan & D.B.Munro (Lamiaceae), all of which, like Erythranthe, belong to the order Lamiales. In Oregon, confirmed hosts also include plants in three additional orders— Asterales (Asteraceae: Petasites frigidus (L.) Fr.), Boraginales (Boraginaceae: Myosotis scorpioides L.), and Caryophyllales (Montiaceae: Claytonia sibirica L.). Larvae and mines have also been photographed on Myosotis in Washington and on Claytonia in British Columbia. Leaf-mining chironomids representing this or a similar species have been found on still other plants in the Pacific Northwest; see the final section of this paper for details. The leaf mines of M. erythranthei are essentially linear throughout, initially narrow and gradually widening, sometimes becoming somewhat irregular or blotchy toward the end. They are mostly full-depth (all of the mesophyll being consumed, leaving only the leaf epidermises intact), with frass in roughly oval pellets that are scattered at random (Figs. 4a, b, d, f). The larvae are able to exit their mines and establish new ones (Figs. 4c, e, g). In a thorough review of the literature on North American leaf-mining insects (Eiseman 2022), we found no records of any leafminer occurring on Erythranthe in nature. The only published record involving this host genus is that of Oatman (1959), who conducted greenhouse experiments with the polyphagous fly Liriomyza sativae Blanchard, 1938 (Agromyzidae). He observed mining in E. guttata; a few larvae survived to the point of pupating, but no adults emerged. Were Liriomyza mines to occur on Erythranthe in nature, they would be easily recognizable by the characteristic pattern of frass in strips on alternating sides of the channel. The only previously known leafminer of Veronica spp. in North America is another agromyzid, Phytomyza crassiseta Zetterstedt, 1860. The formation of the puparium (as opposed to a naked pupa) within the leaf is the most obvious feature distinguishing mines of P. crassiseta from those of M. erythranthei. Claytonia likewise has a single recorded leafminer, the Caryophyllales generalist Pegomya flavifrons (Walker, 1849) (Anthomyiidae). The persistent white eggshell at the beginning and the distinctly blotchy ultimate shape will distinguish mines of P. flavifrons from those of M. erythranthei. The other confirmed hosts of M. erythranthei are also mined by various agromyzid species, but all of these produce mines that are more or less confined to one leaf surface and differ in shape and frass pattern (Eiseman 2022). In the three leaf-mining Cricotopus species, pupation occurs within the mine, but when the adult is ready to emerge, the pupa breaks out of the mine and swims to the water surface (Berg 1950). The habits of M. erythranthei are generally similar. On Erythranthe in British Columbia and on Veronica in Iowa, mines have been collected with pupae inside, each surrounded by a gelatinous mass (Figs. 5a–c, e). In collections of larvae from Claytonia and Erythranthe elsewhere, pupation has taken place outside the mines (at least sometimes in a gelatinous mass), but the leaves had more or less deteriorated by this point. Based on our observations of M. erythranthei on Veronica, the pupa remains in the mine until shortly before emergence of the adult. In one instance the pupal exuviae were found protruding from the mine, with only the tip of the abdomen still inside (Fig. 5d). In another, the exuviae were found loosely adhering to plant material seve, Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 43-51, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Eiseman, C. (2022) Leafminers of North America. Privately published e-book, clxvii + 2213 pp. Available from: http: // charleyeiseman. com / leafminers (accessed 9 January 2023)","Oatman, E. R. (1959) Host range studies of the melon leaf miner, Liriomyza pictella (Thomson) (Diptera: Agromyzidae). Annals of the Entomological Society of America, 52, 739 - 741. https: // doi. org / 10.1093 / aesa / 52.6.739","Berg, C. O. (1950) Biology of certain Chironomidae reared from Potamogeton. Ecological Monographs, 20, 83 - 101. https: // doi. org / 10.2307 / 1943546"]}

Limnophyes spp. (Fig. 7b) Material examined. USA: IOWA: Winneshiek Co., Twin Springs Park, 11.vi.2017, em. ~ 21.vi.2017, J. van der Linden, ex Veronica anagallis-aquatica (2♀); same data but collected 29.i.2018 (1 larva). These specimens were examined by A. Namayandeh in 2018 and their current location is unknown. Biological notes. Each of the collections listed above consisted of a few Veronica stems with attached leaves in which chironomid larvae were forming linear mines. The only larva photographed from the first collection was consistent with M. erythranthei. The two female Limnophyes adults emerged about ten days after the first collection. One of these was light reddish-brown and was identified as Limnophyes cf. carolinensis Saether, 1975 (https:// bugguide.net/node/view/1396485). The other was grayish-black and appeared to be L. viribus (https://bugguide. net/node/view/1396495). The only other insect to emerge from this collection (about a day earlier) was an adult of Scaptomyza pallida (Zetterstedt, 1847) (Drosophilidae), a saprophagous species that sometimes develops as an inquiline in mines made by other fly larvae, including Zygoneura calthella Eiseman, Heller & Rulik, 2016 (Sciaridae) on marsh marigold (Ranunculaceae: Caltha palustris L.) (Eiseman et al. 2016) and Agromyza parvicornis Loew, 1869 (Agromyzidae) on corn (Poaceae: Zea mays L.) (C. Eiseman, unpublished). Whether the Limnophyes adults arose from the observed leaf-mining larvae or were themselves inquilines or contaminants was not known with certainty. About two weeks after the second collection of Veronica leaf mines (on 12 February 2018), actively wriggling Metriocnemus pupae were observed in the leaf mines, and in one case, in a stem. On the same date, an adult male of another orthocladiine, Corynoneura lobata Edwards, 1924, was found dead in the rearing container; its pupal exuviae were not located and the larval habits of this individual are unknown. Corynoneura lobata has an aquatic larva inhabiting mainly fast-flowing waters (Cranston 1982). Also on this date, the now preserved Limnophyes larva, which was initially identified as L. cf. carolinensis but appears consistent with L. viribus, was observed moving within an existing linear mine that was substantially wider than its body, consuming small patches of mesophyll here and there along the margins (Fig. 12b; video: https://youtu.be/0h3Bvc0DfgE). The following day, the first adult Metriocnemus emerged from one of the pupae in the mines. Based on the available evidence, we believe that the Limnophyes species we have reared do not establish their own mines but feed as inquilines in mines created by Metriocnemus larvae., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 61-62, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Eiseman, C. S., Heller, K. & Rulik, B. (2016) A new leaf-mining dark-winged fungus gnat (Diptera: Sciaridae), with notes on other insect associates of marsh marigold (Ranunculaceae: Caltha palustris L.). Proceedings of the Entomological Society of Washington, 118, 519 - 532. https: // doi. org / 10.4289 / 0013 - 8797.118.4.519","Cranston, P. S. (1982) A key to the larvae of the British Orthocladiinae (Chironomidae). Freshwater biological association scientific publication, 45, 1 - 152."]}

Paraphaenocladius exagitans exagitans (Johannsen, 1905) and Paraphaenocladius impensus impensus (Walker, 1856) (Fig. 13) Material examined. USA: IOWA: Allamakee Co., Yellow River State Forest, 43.174434, -91.221148, 13.x.2017, em. 23.xi.2017, J. van der Linden, ex Marchantia polymorpha (1♀, P. exagitans exagitans, ANC); same but em. 2.xii.2017 (1♀, P. exagitans exagitans, USNM); same but em. 1.i.2018 (1♀, P. impensus impensus, USNM); same but em. 15.i.2018 (1Ô, P. exagitans exagitans, ANC); same but em. 29.i.2018 (1♀, P. impensus impensus, ANC); same but em. 12.ii.2018 (1♀, P. exagitans exagitans, USNM); OREGON: Lane Co., Blue River, 44.1535, -122.328, 28.iv.2022, em. 29.iv–7.v.2022, leg. M. W. Palmer, ex Marchantia sp. (2ÔÔ, 1♀, 1 larva, P. exagitans exagitans, USNM; 1Ô, P. exagitans exagitans, ANC). Biological notes. The nominate subspecies of P. exagitans is known from Ontario and throughout the USA as well as China and Japan. It has previously been reared from moist soil at the margins of rivers, streams, seepage, and springs, but none of the published label data mention herbivory (Saether & Wang 1995). Paraphaenocladius impensus s. str. is recorded from Manitoba, Minnesota, Greenland, and northern Europe, where it has been found in moist soil at the margins of lakes, springs, meadows, and alder carr; there have likewise been no observations of herbivory in this species (Saether & Wang 1995). Nematoceran larvae were found mining in thalli of the liverwort Marchantia polymorpha L. (Marchantiaceae) in Iowa in mid-October 2017. The habitat was a ditch in which the soil varied from saturated to submerged in several centimeters of standing water. The larvae initially formed digitate mines with short radiating galleries, and later formed brown blotches. Unlike Metriocnemus mines, which have frass scattered throughout, these mines were mostly clean with frass accumulating around the rim of the entry hole. The first adult of P. exagitans emerged on 23 November from an oblong, dark brown cocoon, composed of frass, which was formed on the surface of a thallus (Figs. 13a, b). Its pupal exuviae were left on the surface of the cocoon. The frass appeared to have been produced by the larva feeding on the surface of the thallus. Two other adults of P. exagitans and three of P. impensus emerged over the next few months. Two larvae were observed mining in fresh thallus growth in late December. Unfortunately, none of the larvae were preserved, and we are unable to confirm that either Paraphaenocladius species feeds as a thallus miner. The photographed larvae are all Sciaridae, and although this rearing effort did not produce any adult sciarids, in 2022 JvdL reared adult sciarids from similar larvae found mining liverwort thalli at another site in Iowa. The sciarid larvae pupated inside their mines. A collection of Marchantia cf. polymorpha in Oregon in late April 2022 produced four adults and a larva of P. exagitans exagitans, along with eight adults of Boreochlus persimilis. Although some empty mines of an unknown insect were seen in a nearby colony of this plant, none were observed in the collected sample, and since the rhizoids of the collected plants were thick and encompassed a substantial amount of mucky organic matter, it is unclear what these midges were feeding on as larvae. The repeated rearing of Paraphaenocladius adults from collections of Marchantia suggests more than a casual association with this plant, but further investigation is required to determine whether the larvae feed as thallus miners, as secondary inhabitants in sciarid mines, as external feeders on the thallus surface or concealed in the rhizoids, or some combination of these., Published as part of Eiseman, Charles S., Namayandeh, Armin, Linden, John Van Der & Palmer, Michael W., 2023, Metriocnemus erythranthei sp. nov. and Limnophyes viribus sp. nov. (Diptera: Chironomidae: Orthocladiinae): leafminers of monkeyflowers, speedwells, and other herbaceous plants, with new observations on the ecology and habitats of other leaf-mining Chironomidae, pp. 41-68 in Zootaxa 5249 (1) on pages 62-64, DOI: 10.11646/zootaxa.5249.1.3, http://zenodo.org/record/7685232, {"references":["Saether, O. A. & Wang, X. (1995) Revision of the genus Paraphaenocladius Thienemann, 1924 of the world (Diptera: Chironomidae, Orthocladiinae). Entomologica scandinavica Supplement, 48, 3 - 69."]}

Updated key to the species of Liriomyza feeding on Euphorbia 1. Mesonotum with yellow longitudinal bands................................................................ 2 1′. Mesonotum largely black, without distinct yellow longitudinal bands............................................ 3 2. Back of head, at level of vertical bristles, entirely yellow. Acrostichal setulae less numerous (5 to 8), irregularly arranged in two rows...................................................................... Liriomyza euphorbiae Martinez [mine initially narrow and serpentine, developing into an irregular blotch; on E. kotschyana Fenzl and E. macroclada Boiss.; Turkey] 2′. Back of head, at level of vertical bristles, partly brown. Acrostichal setulae more numerous, arranged in four almost regular rows...................................................................... Liriomyza heringi Nowakowski [mine a blotch; on E. esula L., E. nicaeensis All., E. palustris L., E. salicifolia Host, and possibly E. amygdaloides L.; Lithuania, Romania, former Soviet Union] 3. Wings without vein dm-cu.............................................................................. 4 3′. Wings with vein dm-cu................................................................................. 5 4. Orbits between upper ors and vti completely yellow. Abdominal tergites without yellow lateral band. Penultimate section of M4 2.5 to 3 times longer than last section.............................................. Liriomyza balcanica (Strobl) [mine broadly linear or a linear-blotch; on E. cyparissias L., E. esula, E. myrsinites L., and possibly E. amygdaloides; Germany and Poland to Turkey] 4′. Upper part of orbits black. At least the basal part of abdominal tergite with yellow border. Penultimate section of M4 maximum 2 times longer than last section................................................... Liriomyza myrsinitae Hering [mine a blotch preceded by a short linear portion; on E. myrsinites; Bulgaria, Spain] 5. Orbits between upper ors and vti completely yellow. Mesophallus long, narrow, fused to distiphallus with broad, bilobed cuplike base from which one pair of short weakly scleritized tubules emerge.................. Liriomyza pascuum (Meigen) [mine a primary blotch, generally containing several larvae; on E. amygdaloides, E. characias L., E. dulcis L., E. esula, E. glareosa Pall. ex M.Bieb., E. palustris, and E. pithyusa L.; throughout Europe] 5′. Upper part of orbits black. Phallus not as above............................................................. 6 6. Frons yellow at base of inner vertical seta. Distiphallus entirely split, short, with minute basal stem contrasting rounded apical chamber...................................................................... Liriomyza strigata (Meigen) [branching mine centered on the midrib; a polyphagous species, uncommonly reported from unspecified Euphorbia; throughout Europe] 6′. Frons brown at base of inner vertical seta. If distiphallus entirely split and short (L. huidobrensis), then apical chamber narrow, not much wider than stem............................................................................... 7 7. Surstylus with two apical spines. Distiphallus angled dorsally, with very short apical chamber........................ 8 7′. Surstylus with one apical spine. Distiphallus various, but not as above........................................... 9 8. Eye 4.8–5.9 times higher than gena. Posterior margin of scutum along scutellum black. Ventral 2/3 of anepisternum brown. Femora entirely yellow. Left distal margin of basiphallus long, extending to apex of paraphallus. Basal stem-like half of distiphallus (fused to mesophallus) gradually narrowing to point of fusion with mesophallus............................................................................................ Liriomyza euphorbiella Eiseman & Lonsdale [mine initially narrow and linear with the frass in alternating strips, expanding to an elongate blotch with irregular and more or less diffuse frass; on E. heterophylla var. cyathophora; USA] 8′. Eye 3.0 times higher than gena. Posterior margin of scutum narrowly yellow. Anepisternum yellow with brown clavate spot. Femora brown basally (paler on venter). Left distal margin of basiphallus shorter, not reaching apex of paraphallus. Basal stemlike half of distiphallus parallel-sided and abruptly wider than mesophallus.. Liriomyza euphorbivora Eiseman & Lonsdale [mine a roughly circular blotch on upper leaf surface, centered on the midrib, with frass in a few small, dark grains and lumps; on E. marginata; USA] 9. Femora usually only brown basally, but sometimes more heavily mottled. Abdominal tergites narrowly yellow laterally. Acrostichal setulae (acr) between second and fourth pairs of dorsocentral setae in four more or less regular rows. Distiphallus simple, cup-shaped................................................................... Liriomyza sativae Blanchard [mine linear; a polyphagous species, with one questionable record from unspecified Euphorbia (light mining of E. pulcherrima observed in a laboratory setting, but no puparia were formed); North and South America; introduced in Europe, Asia, Africa, Australia, and several Pacific islands] 9′. Femora brown basally and with additional light to heavy dorsal mottling. Lateral margin of abdominal tergites brown. Acros- tichal setulae (acr) between second and fourth pairs of dorsocentral setae in two to three more or less regular rows. Distiphallus divided into one pair of short tubules......................................... Liriomyza huidobrensis (Blanchard) [mine linear, often following the midrib and lateral veins; a polyphagous species, uncommonly reported from unspecified Euphorbia according to Martinez & Sobhian (2000); the only record listed by Weintraub et al. (2017) is one of E. marginata from China; this South American fly is introduced in North America, Europe, Asia, and Africa] Characters of the thorax of Liriomyza euphorbivora best differentiate it from other Liriomyza externally, as four rows of acrostichal setulae and a very narrow yellow posterior margin on the scutellum is not found in many species. In the United States, it occurs in L. eupatoriella Spencer, but this species has both vertical setae on yellow, the femora are entirely yellow (not basally brown) and the distiphallus is of a different structure, being more cup-shaped with the base strongly narrowed in lateral view. In Canada, the new species keys most readily to L. cracentis Lonsdale and L. agrios Lonsdale, although the spot on the anepisternum of these two species is much larger, their femora are entirely yellow, and the distiphallus is more cup-shaped, without a short apical chamber. The phallus instead shows more similarity to species without the posteriorly yellow scutum. These include L. conclavis Lonsdale (host unknown), L. asclepiadis Spencer and L. peleensis Spencer (both on Asclepias), which have a medially constricted mesophallus and a very sharply upturned distiphallus, and L. subasclepiadis Spencer (also on Asclepias), which also has a more strongly angled distiphallus., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 28-29, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Martinez, M. & Sobhian, R. (2000) Review of the agromyzid flies associated with Euphorbia spp., and prospects of their use for biological control of leafy spurge in North America. Redia, 81, 1 - 15. [1998]","Weintraub, P. G., Scheffer, S. J., Visser, D., Valladares, G., Correa, A. S., Shepard, B. M., Rauf, A., Murphy, S. T., Mujica, N., MacVean, C., Kroschel, J., Kishinevsky, M., Joshi, R. C., Johansen, N. S., Hallett, R. H., Civelek, H. S., Chen, B. & Metzler, H. B. (2017) The invasive Liriomyza huidobrensis (Diptera: Agromyzidae): understanding its pest status and management globally. Journal of Insect Science, 17 (1), 1 - 27. https: // doi. org / 10.1093 / jisesa / iew 121"]}

Ophiomyia abutilivora Spencer (Figs. 64–65) Material examined. IOWA: Winneshiek Co., Decorah, Trout Run Trail, vii.2017, em. by 26.vii.2017, J. van der Linden, ex Abutilon theophrasti, # CSE4940, CNC1643668–1643669 (1♁ 1♀). Hosts. Malvaceae: Abutilon theophrasti Medik.; adults have been collected on flowers of Sida cordifolia L. (Spencer & Steyskal 1986). Larval mine. (Figs. 64–65) The larva mines up and down the stem, forming an irregular, gall-like “welt” and associated scarring. Puparium. Whitish to yellowish; formed within the stem, with the anterior spiracles projecting through the epidermis. Phenology and voltinism. The only reared specimens in the type series are from Mississippi with a date of 22 October; whether this was the collection or emergence date was not stated, but either way they would seem to represent a different generation from our Iowa specimens, which emerged from green stems in late July. Adults have been collected in August and September in Minnesota, and in March in Florida (Spencer & Steyskal 1986). Distribution. USA: FL, *IA, [IL], MN, MS, [WI] (Spencer & Steyskal 1986)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 16, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Phytomyza flavicornis Fallén (Fig. 57) Material examined. IOWA: Winneshiek Co., Roslien Woods, 1.ii.2018, em. 18.ii.2018, J. van der Linden, ex Urtica dioica, # CSE4943, CNC1643673–1643674 (1♁ 1♀). Hosts. Urticaceae: Urtica dioica L.; adults have been caught on U. gracilis Aiton (Spencer 1969). Larval biology. The larva bores in the pith of the stem and roots. Puparium. (Fig. 57) Slender and yellowish, with deep intersegmental grooves; formed within the stem. Phenology and voltinism. Pupae overwinter, with adults emerging in spring. Collection dates for adults range from 23 April in Oregon to 7 July in Manitoba (Spencer 1969; Spencer & Steyskal 1986). Distribution. USA: *IA, IN, MI, OR; Canada: AB, MB, ON; Europe (Spencer 1969; Spencer & Steyskal 1986)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 34, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. (1969) The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada, 64, 1 - 311. https: // doi. org / 10.4039 / entm 10164 fv","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Phytomyza solidaginophaga Sehgal Material examined. IOWA: Winneshiek Co., Decorah, Twin Springs Park, 24.v.2017, em.?, J. van der Linden, ex Solidago flexicaulis, # CSE4670, CNC1144022 (1♁). Hosts. Asteraceae: Solidago caesia L., S. canadensis L., S. * flexicaulis L., S. patula Muhl. ex Willd., S. rugosa Mill., S. simplex Kunth (Eiseman & Lonsdale 2018); see Comments for other likely hosts. Leaf mine. Greenish-white to greenish-brown, entirely linear, with frass in closely spaced grains, partly forming short beaded strips (Eiseman & Lonsdale 2018). Puparium. Black; formed outside the mine. Phenology and voltinism. This species is univoltine, overwintering as pupae, with larvae present only in spring. Although Alberta collection dates range from 7 to 13 June (Griffiths 1976), larvae have only been found from 11 to 24 May in the USA (Eiseman & Lonsdale 2018). Distribution. USA: CO (Winkler et al. 2009), *IA, MA; Canada: AB, [YT] (Eiseman & Lonsdale 2018). Comments. Given that the latest recorded date for larvae of Phytomyza solidaginophaga in the USA (24 May) is more than two weeks before the earliest recorded date for P. astotinensis (9 June; this paper), we believe mines found in Massachusetts on two additional hosts likely represent P. solidaginophaga: occupied mines on Solidago gigantea Aiton between 18 and 22 May, and a mine on S. arguta Aiton containing a dead larva on 25 May. An occupied mine on S. speciosa Nutt. photographed on 31 May in Wisconsin (Klein 2019) may also represent P. solidaginophaga., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 37-38, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Griffiths, G. C. D. (1976) Studies on boreal Agromyzidae (Diptera). XII. Phytomyza and Chromatomyia miners on Astereae (Compositae). Quaestiones entomologicae, 12, 239 - 278.","Winkler, I. S., Scheffer, S. J. & Mitter, C. (2009) Molecular phylogeny and systematics of leaf-mining flies (Diptera: Agromyzidae): delimitation of Phytomyza Fallen sensu lato and included species groups, with new insights on morphological and host-use evolution. Systematic Entomology, 34, 260 - 292. [including online Supporting Information] https: // doi. org / 10.1111 / j. 1365 - 3113.2008.00462. x","Klein, T. (2019) Leaf mine. Available from: https: // bugguide. net / node / view / 1668919 (accessed 15 December 2019)"]}

Agromyza aristata Malloch (Fig. 59) Material examined. NORTH CAROLINA: Scotland Co., Laurinburg, St. Andrews University, 18.iv.2017, em. ~ 17.iv.2018, T. S. Feldman, ex Celtis laevigata, # CSE4418, CNC1144090 (1♀); 19.iv.2017, em. ~ 17.iv.2018, T. S. Feldman, ex Celtis laevigata, # CSE4417, CNC1135675 (1♀); OKLAHOMA: Payne Co., Mehan, 36.013839°, - 96.998103°, 7.v.2017, em. 24.iv.2018, M.W. Palmer, ex Celtis occidentalis, # CSE4435, CNC1135641 (1♁). Photographed mines. ALABAMA: Madison Co., Owens Cross Roads, 2.v.2020, kidneymoth, Ulmus alata [vacated mines], iNat 44595725; Redstone Arsenal, 18.iv.2020, kidneymoth, Ulmus americana [vacated mine], iNat 42532868; 19.v.2020, kidneymoth, Ulmus rubra [vacated mine], iNat 46547423; COLORADO: Boulder Co., Boulder, S 45 th St., 14.vii.2019, J. Shorma, Ulmus [vacated mine], iNat 32754344; CONNECTICUT: Hartford Co., Windsor, 41.893658, -72.634136, 29.v.2020, M. Apgar, Ulmus, iNat 47765803; GEORGIA: Fulton Co., Atlanta, Simmons Lake, 2.v.2020, E. Summerbell, Ulmus americana [vacated mine], iNat 44678255; KANSAS: Brown Co., Horton, 25.v.2020, ksgardener1, Ulmus [vacated mine], iNat 47359824; MASSACHUSETTS: Franklin Co., Gill, Pisgah Mountain Rd., 29.iv.2012, C.S. Eiseman, Ulmus; MARYLAND: Baltimore City County, Herring Run Park, 6.v.2015, T. Wilson, Ulmus, BG 1067957; MICHIGAN: Marquette Co., Ives Lake, 20.vi.2019, C.S. Eiseman, Ulmus [vacated mine]; MINNESOTA: Scott Co., Prior Lake, Cleary Lake Regional Park, 44.690462, -93.391813, 16.vii.2019, A. Manning, Ulmus rubra [vacated mines], iNat 27098197; NEW JERSEY: Somerset Co., Warren Twp., 10.v.2011, S. Rall, Ulmus, BG 514546; NORTH CAROLINA: Durham Co., Durham, Stagecoach Road Eagle Spur Trail, 10.vii.2017, T.S. Feldman, Ulmus alata [vacated/predated mine], BG 1401462; Wake Co., Lake Crabtree County Park, 27.vi.2018, T.S. Feldman, Ulmus alata [vacated/predated mine], BG 1763158; NORTH DAKOTA: Burleigh Co., Bismarck, 804 N Third St., 20.vi.2020, megnd, Ulmus [vacated mines], iNat 50350205; TENNESSEE: Chester Co., Henderson, 27.iv.2015, K. Childs, Ulmus [mine with dead larva], BG 1179873; WIS- CONSIN: Dane Co., Cross Plains, 28.v.2011, I. Loser, Ulmus [aborted mine], BG 614335; TEXAS: Harris Co., Houston, Wedgehill Ln., 24.iv.2020, D. Johnson, Celtis pallida [vacated/aborted mine], iNat 43187133; Tarrant Co., Keller, 10.iv.2020, K. Sasan, Ulmus americana, iNat 42888081; Tarrant, 7.iv.2020, K. Sasan, Ulmus americana, iNat 41740783; NEW BRUNSWICK: Sunbury Co., Maugerville, 17.vi.2015, C. Adam, Ulmus [mine with dead larva], BG 1162296; QUÉBEC: Communauté-Urbaine-de-Québec, Neufchâtel Est–Lebourgneuf, 5.vi.2020, C. Grenier, Ulmus [vacated/aborted mine], iNat 48760929. Hosts. * Cannabaceae: [Celtis laevigata Willd.], C. occidentalis L., [C. pallida Torr.]; Ulmaceae: [Ulmus alata Michx.], U. americana L., U. rubra Muhl. (Eiseman & Lonsdale 2018). Leaf mine. (Fig. 59) On Ulmus, the mine is formed on the upper surface, usually beginning near the leaf margin; initially green and linear, later widening into a brown, elongate blotch. Frass is indistinct in the narrow, green portion, but later present in blackish particles that often form two rows, becoming more randomly scattered toward the end. Concentric feeding lines are evident in the blotch (Eiseman & Lonsdale 2018). Mines on Celtis (Fig. 59) are more often formed on the lower leaf surface, may begin well away from the leaf margin, and tend to remain essentially linear throughout their length, though sometimes contorted so that this is not obvious. Puparium. Yellowish-brown to reddish-brown; formed outside the mine. Phenology and voltinism. This species is univoltine, with larvae feeding in April and May; pupae overwinter. Distribution. USA: [AL, AR, CO, CT, FL, GA], IL, IN, IA, [KS, MA, MD, MI, MN, *NC, ND, NJ], NY, OH, OK, PA, [TN], VA, VT, [WI, TX]; Canada: AB, [NB], ON, [QC] (Eiseman & Lonsdale 2018). Comments. Liriomyza adults were reportedly reared from the mines on Ulmus americana in Keller, TX that are cited above (Sasan 2020). The mines were collected on 10 April, puparia appeared on 13 April and were transferred to glass jars, and adults emerged on 21 April. Repetition of this rearing would be desirable; we suspect that a mixup of puparia with some from another rearing occurred, since the photographed mines are identical with those of Agromyza aristata. Five closely related Agromyza species are now known to feed on Celtis. The entirely yellow legs easily distinguish A. aristata from the others— A. fission Eiseman & Lonsdale, A. varifrons Coquillett, the gall inducer A. deserta (Patton), and the European A. trebinjensis Strobl—which all have mostly dark legs (Strobl 1900; Spencer & Steyskal 1986; von Tschirnhaus 2017; Eiseman & Lonsdale 2018). Whereas in A. fission, A. trebinjensis, and A. varifrons the posterior spiracles of the larva each have three bulbs as is typical of the genus, those of A. aristata have 8–10, and those of A. deserta over 20., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 5-6, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Sasan, K. (2020) [Photographed leaf mines, puparia, and reared flies.] Available from: https: // www. inaturalist. org / observations / 42891605 (accessed 15 July 2020)","Strobl, P. G. (1900) Dipterenfauna von Bosnien, Hercegovina und Dalmatien. Wissenschaftliche Mittheilungen aus Bosnien und der Hercegovina, 7, 552 - 670.","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","von Tschirnhaus, M. (2017) First description of the gall fly Agromyza deserta (Patton) comb. nov. (Diptera: Agromyzidae), originally classified in an ichnogenus and now reared from Celtis occidentalis Linnaeus (Cannabaceae). Studia dipterologica, 23 (1), 151 - 161. [2016]"]}

Liriomyza taraxaci Hering (Fig. 79) Material examined. VERMONT: Washington Co., Montpelier, North Branch River Park (44.283214, -72.571026), 21.vii.2018, em. 9.viii.2018, C.S. Eiseman & J.A. Blyth, ex Taraxacum officinale, # CSE4914, CNC1643637 (1♀); ONTARIO: Renfrew Co., Pembroke, Pansy Patch Park, 45.821464, -77.112117, 7.vii.2018, em. 20.vii.2018, C.S. Eiseman & J.A. Blyth, ex Taraxacum officinale, # CSE4811, CNC1643630 (1♁). Tentatively identified material. NORTH CAROLINA: Scotland Co., Laurinburg, St. Andrews University, 10.iv.2017, em. 6.v.2018, T. S. Feldman, ex Krigia virginica, # CSE4499, CNC1144028 (1♀). Photographed mines. MARYLAND: Baltimore City Co., Herring Run Watershed, 19.vi.2017, T. Wilson, Taraxacum? officinale, BG 1228804; MINNESOTA: Fillmore Co., Rushford, Magelssen Bluff Park, 24.vi.2019, C.S. Eiseman, Lactuca canadensis [vacated mines]; NORTH CAROLINA: Wake Co., Lake Crabtree County Park, 9.v.2019, T.S. Feldman, Krigia dandelion [vacated mines], BG 1660443. Hosts. Asteraceae: [Cichorium intybus L., Krigia dandelion Nutt., K. * virginica (L.) Willd.], Lactuca biennis (Moench) Fernald, L. canadensis L., L. sativa L., [L. serriola L.], Taraxacum officinale F.H.Wigg. (Eiseman & Lonsdale 2019). Leaf mine. (Fig. 79) The mines on Krigia are linear throughout, necessarily contorted in the small leaves, with frass in grains, irregular particles, and short strips along the sides. Mines of L. taraxaci s.l. on Lactuca and Taraxacum rapidly widen to form blotches, and larvae are frequently gregarious on Lactuca (Eiseman & Lonsdale 2018, 2019). Puparium. Yellow to yellow-orange; formed outside the mine. Phenology and voltinism. The tentatively identified female was collected as a larva in early April and the adult emerged the following spring. Otherwise, adults we have reared of Liriomyza taraxaci s.l. have emerged 11–25 days after the larval collection dates, indicating the species is at least bivoltine. These have included larvae collected in North Carolina on 28 April and 10 June, and in Massachusetts on 6 July, 12 July, and 21 August (Eiseman & Lonsdale 2018, 2019). In Alberta, a larva collected in August apparently emerged as an adult the following spring, unless the error in the reported emergence date was in the month rather than the year (collected 18.viii.1974, emerged 5.v.1974, according to Lonsdale (2017a)). Distribution. USA: AK, IL, MA, [MD, MN], NC, NY, *VT, WA, [WI]; Canada: AB, BC, MB, NB, ON, QC, SK, YT; Europe (Eiseman & Lonsdale 2019). Comments. The only previous record of an agromyzid from Krigia is that of Ophiomyia beckeri (Hendel), also in North Carolina (Eiseman et al. 2019). The year-long diapause observed in this rearing of Liriomyza cf. taraxaci from Krigia is highly unusual for Liriomyza in our experience. Due to this, along with the entirely linear leaf mines, the rearing of males to confirm the identity of the Krigia feeders is desirable. The Taraxacum officinale -reared specimens in collections CSE4811 and CSE4914 reveal previously unappreciated variation in Liriomyza taraxaci, as both have a very narrow yellow line along the posterior margin of the scutum. This character will key it to L. cracentis in Lonsdale (2017a), but they differ in that the posterior yellow margin is much narrower (almost indistinguishable) and doesn’t narrowly extend anteriorly at two points to touch the bases of the posterior pair of dorsocentrals. Furthermore, the bases of the femora are brown dorsally and the fore femur has brownish streaking, as seen in L. taraxaci. We have stated that L. taraxaci is a relatively variable species, so this additional small variation is not entirely surprising if it is conspecific. Material discussed in Lonsdale (2017a) was re-examined, and this line is present in some specimens collected in Ottawa., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 31, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Lonsdale, O. (2017 a) The Liriomyza (Agromyzidae: Schizophora: Diptera) of Canada & Alaska. Zootaxa, 4234 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4234.1.1"]}

Ophiomyia cf. frosti Spencer (Fig. 69) Material examined. OKLAHOMA: Payne Co., Meham, 36.014339° N, - 96.996744° W, 16– 21.iii.2018, em. 29.iii.2018, M.W. Palmer, ex Lactuca, # CSE4412, CNC1135643 (1♀); 23.iii.2018, em. 8.iv.2018, # CSE4413, CNC1135644–1135646 (3♁); 1.iv.2018, em. 8.iv.2018, # CSE4414, CNC1144112–1144113 (2♀); 4.iv.2018, em. 15.iv.2018, # CSE4453, CNC1135650 (1♁). Host. * Asteraceae: Lactuca L. Leaf mine. (Fig. 69) Whitish, upper-surface (nearly full-depth), extending along the midrib from the base of the petiole to the blade, where irregular lobes or branches extend to either side (these may be partially green and interparenchymal); ~ 5 cm long in total. Frass is deposited densely along the midrib. Puparium. Black, formed within a leaf (not necessarily the one with the conspicuous mine), on the lower surface near the base of the petiole, with the anterior spiracles projecting through the lower epidermis. Phenology and voltinism. In Oklahoma, larvae and puparia (some already empty) have been found beginning in mid-March, with adults emerging from late March to mid-April. The holotype of O. frosti was collected in New York on 16 May (Spencer & Steyskal 1986). Distribution. USA: NY (Spencer & Steyskal 1986), *OK. Comments. Ophiomyia frosti was described from a single caught adult. Spencer & Steyskal (1986) noted that it closely resembles O. pulicaria (Meigen), known from western Canada and common in Europe, where it feeds on various genera of Cichorieae including Lactuca, forming midrib-based mines that are apparently indistinguishable from those from which our Oklahoma specimens were reared. The new specimens differ from the type in usually having two ori (not three), but there is sometimes one on one side, and one male has three on one side. The specimens are slightly larger, with a wing length of 2.8–3.0 mm (not 2.3 mm); the male genitalia appear to match those of the holotype (Spencer & Steyskal 1986: Figs. 187, 188). Ophiomyia pulicaria differs in being smaller (wing length 1.9–2.2 mm), and in having a very broad and apically flattened distiphallus and an H-shaped basiphallus (ventral view—see Spencer (1976: Fig. 97))., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 17, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Agromyza reptans Fallén Material examined. MASSACHUSETTS: Franklin Co., Northfield, 276 Old Wendell Rd., 22.x.2017, em. 30.iv.2018, C.S. Eiseman, ex Urtica dioica, # CSE4459, CNC1144082 (1♀). Host. Urticaceae: Urtica dioica L. Leaf mine. Blackish, with paler greenish-brown patches only in the newest portions where the larvae are actively feeding; initially narrow and following the leaf margin, later expanding to a broad blotch. Frass is partly diffuse and partly in irregular lumps. Puparium. Reddish-brown; formed outside the mine. Phenology and voltinism. This species has at least two generations per year; it has not been observed to have a summer pupal diapause as observed in A. pseudoreptans, but there is evidently some variation in length of the pupal stage. Spencer (1969) listed a specimen emerging on 14 August from a larva collected on 5 June in Ontario, whereas Eiseman & Lonsdale (2018) listed specimens emerging on 21 July from larvae collected on 28 June in New York. The larvae from which our Massachusetts specimen was reared were found on 22 October, a few days before larvae of A. pseudoreptans appeared several meters away. Distribution. USA: CA, *MA, NY, WA; Canada: ON, QC, SK; Europe; Kazakhstan; Turkmenistan (Nartshuk & von Tschirnhaus 2017; Eiseman & Lonsdale 2018)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 7, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. (1969) The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada, 64, 1 - 311. https: // doi. org / 10.4039 / entm 10164 fv","Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Nartshuk, E. & von Tschirnhaus, M. (2017) Leafminer flies (Diptera: Agromyzidae) of the fauna of Russia and adjacent countries: The genus Agromyza Fallen. Studia dipterologica, 22 (2), 215 - 232. [2015]"]}

Ophiomyia cf. chondrillae Spencer Material examined. IOWA: Allamakee Co., 5.ix.2017, em. spring 2018, J. van der Linden, ex Lactuca, # CSE4671, CNC1135659–1135660 (1♁ 1♀). Hosts. Asteraceae: Chondrilla juncea L. (Spencer & Steyskal 1986), * Lactuca L. Larval mine. An externally visible stem mine. Puparium. Details not recorded by either Spencer & Steyskal (1986) or us, but we believe the puparium was not covered by a “scab” as described in the Comments under Melanagromyza sp. 2. Phenology and voltinism. The holotype of O. chondrillae, from Washington, is labeled with a date of 11 July; whether this was the collection or emergence date was not stated, but there are evidently at least two generations since our Iowa specimens emerged in spring from stems collected in early September. Distribution. USA: *IA, WA (Spencer & Steyskal 1986). Comments. Ophiomyia chondrillae was described from a single specimen in poor condition (Spencer & Steyskal 1986), which we have not examined, but our specimens key readily to this species in Spencer & Steyskal (1986). The morphology of the flies reared from Lactuca seems to fit this species, and we suspect that slight differences in genitalic morphology compared to the original illustrations are due to error or a difference in the angle in which the phallus was illustrated. Given this, and the fact that the host is in the same tribe (Cichorieae) as that of the holotype, we choose to regard them as conspecific until we can examine specimens reared from Chondrilla., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 17, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Melanagromyza arnoglossi Eiseman & Lonsdale, spec. nov. (Figs. 1–4, 60, 87–92) Holotype. USA. IOWA: Winneshiek Co., Chattahoochie Park, 6.xi.2017, em. spring 2018, J. van der Linden, ex Arnoglossum, # CSE4643, CNC1144035 (1♁). Etymology. The specific epithet refers to the host plant genus, Arnoglossum Raf. Host. Asteraceae: Arnoglossum Raf. Larval biology. (Fig. 60) The larva forms a narrow linear gallery (not externally visible) in the thin layer of pith lining the inside of the hollow stem. Puparium. (Fig. 4) Straw-colored; formed within the stem, at the end of the larval gallery. Phenology and voltinism. Pupae overwinter; further details unknown. Distribution. USA: IA. Adult description. Wing length approximately 2.8 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 0.7. Eye height divided by gena height: 4.0. First flagellomere small, round- ed. Orbital plate and dorsal half of parafacial strongly projecting and broadly rounded. Ocellar triangle ill-defined apically where it takes on texture of more rugose vitta, but it likely ends short of lunule. Lunule with medial groove, extending almost to level of second fronto-orbital. Gena somewhat straight with slight upwards curve on anterior half. Cheek wide, not well-defined. Clypeus narrowly rounded anteriorly. Eye and head longest above midpoint. Thorax subshining. Specimen in relatively poor condition, with body partially collapsed, and aristae, fore tarsi, right mid leg and most setae missing. Chaetotaxy: Fronto-orbital setae mostly broken or missing; five setae evident from sockets or remaining bases of setae, possibly becoming shorter and more slender anteriorly, evenly spaced with anterior seta slightly displaced forward. Orbital setulae in several scattered rows; short, appearing erect and lateroclinate. Postvertical seta subequal to ocellar seta. Eye with relatively narrow patch of hairs dorsomedially that are longer than orbital setulae. Most thoracic setae missing dorsally, including dorsocentrals, but two large sockets evident. Acrostichal setulae in ten irregular rows. Katepisternum with two additional subdominant setae. Mid tibia with one posteromedial seta. Coloration: (Figs. 1–3) Setae dark brown. Color dark brown, including halter; gena and anteroventral region of parafacial paler brown; abdomen and thorax green metallic, with color less evident on thorax below notum; wing veins brown. Calypter margin and hairs yellow. Genitalia: (Figs. 87–92) Epandrium with small anteroventral spine. Surstylus fused to anteroventral margin of epandrium, wide and shallow, with two irregular rows of tubercle-like setae along inner surface. Cercus large, welldeveloped. Hypandrium subtriangular, broad basally with thick arms, with narrow tapered apex; inner lobe split, sinuate. Phallophorus cylindrical, base narrow, venter bulging. Basiphallus U-shaped, adjacent to phallophorus, dorsal surface long; distance between it and distiphallus+mesophallus as long as phallophorus. Mesophallus small, cylindrical, fused to ventromedial surface of distiphallus; base of mesophallus and distiphallus level. Distiphallus with one pair of tubules flanking mesophallus; long dorsomedial process enclosed ventrally by narrow plate; dorsal chamber relatively shallow, flat, with one pair of spinulose pads on inner surface; subovate in ventral view, length twice width. Ejaculatory apodeme with narrow base and stem; blade large, pale and ovate with sclerotized medial rib; stem with globose lateral process; sperm pump with transverse sclerotized bar curved up at ends and with long faint tubule emerging from end opposite duct. Comments. This is the first rearing record of any agromyzid from Arnoglossum. The host was a hollowstemmed woodland species; in Iowa we have also found stems of the prairie species A. plantagineum Raf. with the solid pith riddled with galleries of an unknown Melanagromyza species (the empty puparia were similarly strawcolored). In Wisconsin, Klein (2013) photographed what appears to be a vacated Liriomyza mine (with frass in alternating strips) on the lower surface of a leaf of A. atriplicifolium (L.) H.Rob. This dark metallic green species has a frons that strongly projects above the eye, a modestly well-developed ocellar triangle and five fronto-orbitals. This list of features will key it to Melanagromyza cirsiophila in Spencer & Steyskal (1986), which is known on Cirsium and Carduus (Asteraceae) in California and Costa Rica (Shi & Gaimari 2015). The distiphallus of M. cirsiophila differs, however, in being adjacent to the basiphallus, and it is much longer with a broad base and medial constriction. The species M. angelicae (Frost), known from Angelica (Apiaceae) in New York and Ohio, has an additional sixth fronto-orbital and is otherwise quite similar externally. The phallus is also similar, but the length of the distiphallus is equal to the length between it and the basiphallus (not the phallophorus), the distiphallus does not have a pronounced tubule emerging anterodorsally, the dorsal chamber of the distiphallus is highest apically (not basally), and the basiphallus is narrower. Illustrations in Steyskal (1980b) also show the surstylus to be more pronounced and the distiphallus (ventral view) to be shorter compared to its width and more sharply pointed., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 7-8, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Klein, T. (2013) Leaf mine. Available from: https: // bugguide. net / node / view / 817850 (accessed 10 December 2019)","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","Shi, L. & Gaimari, S. D. (2015) Revision of the genus Melanagromyza in California, with descriptions of three new species (Diptera: Agromyzidae). Zootaxa, 4005 (1), 1 - 102. https: // doi. org / 10.11646 / zootaxa. 4005.1.1","Steyskal, G. C. (1980 b) Six new North American species of Melanagromyza (Diptera, Agromyzidae). Journal of the Washington Academy of Science, 70 (1), 36 - 43."]}

Liriomyza eupatorii (Kaltenbach) Material examined. ILLINOIS: Cook Co., Glencoe, Chicago Botanic Gardens, 42°8′56.66″N 87°47′21.99″W, oak woodland, 21.vi.2018, em. by 25.vi.2018, J.F. Steffen, ex Symphyotrichum shortii, CNC 1135573–1135575 (2♁ 1♀); ONTARIO: Nipissing, Algonquin Provincial Park, Basin Depot, 6.vii.2018, em. 16.vii.2018, C.S. Eiseman & J.A. Blyth, ex Symphyotrichum? cordifolium, # CSE 4778, CNC 1643629 (1♁). Hosts. Apocynaceae: Asclepias L.; Asteraceae: Baccharis halimifolia L., Mikania micrantha Kunth, M. scandens (L.) Willd., Solidago altissima L., S. canadensis L., S. latissimifolia Mill., Symphyotrichum chilense (Nees) G.L.Nesom, S. cordifolium (L.) G.L.Nesom, [S. lateriflorum (L.) Á.L̂ve & D.L̂ve, S. novae-angliae (L.) G.L.Nesom], S. praealtum (Poir.) G.L.Nesom, S. puniceum (L.) Á.L̂ve & D.L̂ve, S. * shortii (Lindl.) G.L.Nesom, Xanthium strumarium L. (Eiseman & Lonsdale 2018). California specimens apparently reared from Callistephus Cass. (Asteraceae) have been tentatively identified as L. eupatorii (Lonsdale 2011); see Comments for other likely hosts. Leaf mine. Whitish, upper surface; long, narrow and linear; often beginning with a distinctive spiral, but this is sometimes reduced to a minute contorted area, and sometimes there is no hint of it whatsoever. The black frass is mostly in distinct, alternating strips, sometimes devolving into irregular, squiggly fragments toward the end (Eiseman & Lonsdale 2018). Puparium. Yellow to orange to dark brown; formed outside the mine (Eiseman & Lonsdale 2018). Phenology and voltinism. This species is evidently multivoltine in the USA, although all of our rearing records involve larval collections and adult emergences prior to midsummer. We have reared adults of Liriomyza eupatorii from larvae found in Oklahoma as early as 23 March, and from larvae found in Massachusetts on 11 May, 11–13 June, and 24–28 June. In New York, Scheffer & Lonsdale (2018) similarly recorded larvae only in May and June. Later dates have been observed in Canada, including an Alberta specimen emerging on 19 August from a larva collected on 4 August (Lonsdale 2017a). In all of our rearings, adults have emerged 10–22 days after the larvae were collected. Distribution. USA: CA, DE, GA, *IL, MA, MI, MS, MT, NC, NY, OK, PA, SC, TN, VA, WA, WV; Canada: AB, BC, MB, NB, NS, ON, QC, SK; Europe (Eiseman & Lonsdale 2018). Comments. We have found Liriomyza mines beginning with distinct spirals, and thus likely representing L. eupatorii, on the following additional asteraceous hosts: Ambrosia artemisiifolia L. (MA), Erigeron canadensis L. (MA, NC), Euthamia graminifolia (L.) Nutt. (MA), Senecio triangularis Hook. (ID), Solidago rigida L. (MA), S. sempervirens L. (NY), S. simplex var. randii (Porter) Kartesz & Gandhi (NH), Symphyotrichum firmum (Nees) G.L.Nesom (IL), S. lanceolatum (Willd.) G.L.Nesom (MA), S. pilosum (Willd.) G.L.Nesom (NC), S. ulmifolia Muhl. ex Willd. (OK), and Zinnia elegans Jacq. (NC). Liriomyza mines without spirals, but nonetheless probably representing L. eupatorii, have been found on Solidago patula Muhl. ex Willd. (MA). Papp & Černý (2017) considered Liriomyza eupatorii to be synonymous with the senior L. pusilla (Meigen), which is supported by close overall external and genitalic morphological similarity, notably including general phallic morphology, a large ejaculatory apodeme with a narrowed stem, and the occasional presence of a third ori. Liriomyza pusilla is known from Arctium L., Aster L., Bellis L., Bidens L., Callistephus Cass., Crassocephalum Moench, Epaltes Cass., Hypochaeris L., Synedrella Gaertn., Tithonia Desf. ex Juss., Vernonia Schreb., Solidago L., and Xanthium L. (Asteraceae) (Benavent-Corai et al. 2005; von Tschirnhaus & Karimpour 2006; Ellis 2020), the last two of which are also host genera of L. eupatorii. Beyond its Nearctic hosts, L. eupatorii has been reported from Aster, Eupatorium L., Helianthus L., Lapsana L. (Asteraceae), and Galeopsis L. (Lamiaceae) (Spencer 1976; Benavent-Corai et al. 2005). This synonymy follows notes of similarity by previous authors including Spencer (1971), who stated that adult morphology appeared identical, both external and genitalic, but that the form of the leaf mines was distinct. Mines of L. eupatorii typically start with a characteristic spiral, but we have observed some to lack this spiral (Eiseman & Lonsdale 2018), as seen in L. pusilla. Spencer (1976) further differentiated the two by L. pusilla having the “mesonotum brilliantly shining black” and L. eupatorii having the “mesonotum deep black but less shining”. While these two species are clearly closely related, their synonymy may be unwarranted on the basis of minimal overlap in host genera, a relatively (but not entirely) consistent difference in larval mine pattern, and slight differences in the phallus. While no specimens of L. pusilla were available to us, descriptions and illustrations were examined from Spencer (1971) and Papp & Černý (2017). Lonsdale (2017a) noted variation in the shape of the phallus of L. eupatorii, but in all cases, the mesophallus is quite small, being narrow, stem-like, and no more than half the length of the larger distiphallus, while it is slightly longer in L. pusilla and much thicker in lateral view. Regarding the distiphallus itself, it is longer and usually widest at or past the midpoint of the segment in L. eupatorii, and shorter, rounder, and either parallel-sided or widest on the basal half in L. pusilla (also one atypical male of L. eupatorii from Alberta (Lonsdale 2017a: Fig. 267)). Based on all of the above evidence, the two species are here considered separate (with L. eupatorii being Holarctic and L. pusilla known only from the Palearctic and Oriental regions), but because of their strong similarities, further scrutiny is certainly warranted, ideally including molecular data sets., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 25-26, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Lonsdale, O. (2011) The Liriomyza (Agromyzidae: Schizophora: Diptera) of California. Zootaxa, 2850 (1), 1 - 123. https: // doi. org / 10.11646 / zootaxa. 2850.1.1","Scheffer, S. J. & Lonsdale, O. (2018) A survey of Agromyzidae (Diptera) reared from leafmines on Long Island, New York; host associations, distribution data, and the description and host association of a new species. Zootaxa, 4450 (1), 77 - 90. https: // doi. org / 10.11646 / zootaxa. 4450.1.5","Lonsdale, O. (2017 a) The Liriomyza (Agromyzidae: Schizophora: Diptera) of Canada & Alaska. Zootaxa, 4234 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4234.1.1","Papp, L. & Cerny, M. (2017) Agromyzidae (Diptera) of Hungary. Vol. 3. Phytomyzinae II. Pars Ltd., Nagykovacsi, 427 pp. https: // doi. org / 10.18655 / Agromyzidae. Vol. 3","Benavent-Corai, J., Martinez, M. & Jimenez Peydro, R. (2005) Catalogue of the host plants of the world Agromyzidae (Diptera). Bollettino di Zoologia agraria e di Bachicoltura, Series II, 37 (Supplementum), 1 - 97.","von Tschirnhaus, M. & Karimpour, Y. (2006) First record of Liriomyza pusilla (Meigen, 1830) (Diptera: Agromyzidae) from Iran. Proceedings of the 17 th Iranian Plant Protection Congress, 17 (1), 76.","Ellis, W. N. (2020) Leafminers and plant galls of Europe. Available from: https: // bladmineerders. nl / (accessed 12 September 2020)","Spencer, K. A. (1971) Notes on a revision of the British Agromyzidae (Diptera) including the description of 14 new species. Entomologist's Gazette, 22 (3), 141 - 195."]}

Agromyza parca Spencer Material examined. IOWA: Allamakee Co., Footbridge Farm, 22.vii.2018, em. by 14.viii.2018, J. van der Linden, ex Poaceae, # CSE4946, CNC1643675–1643677 (2♁ 1♀); NORTH CAROLINA: Durham Co., Durham, 17- Acre Wood Preserve, 8.v.2017, em. 4.v.2018, T.S. Feldman, ex Dichanthelium, # CSE4482, CNC1135677–1135678 (1♁ 1♀); Scotland Co., Laurinburg, St. Andrews University, 10.v.2017, em. 6–14.v.2018, T.S. Feldman, ex Dichanthelium, # CSE4502, CNC1144099–1144100 (1♁ 1♀); Wake Co., Morrisville, Lake Crabtree County Park, 6.vi.2018, em. 25.vi.2018, T.S. Feldman, ex Dichanthelium scoparium, # CSE4695, CNC1135686 (1♁). Tentatively identified material. NORTH CAROLINA: Scotland Co., Laurinburg, St. Andrews University, 3.v.2017, em. 6.v.2018, T. S. Feldman, ex Dichanthelium, # CSE4503, CNC1144101–1144102 (2♀). Hosts. Poaceae: Dichanthelium clandestinum (L.) Gould, D. * scoparium (Lam.) Gould, Glyceria canadensis (Michx.) Trin., G. striata (Lam.) Hitchc. (Eiseman & Lonsdale 2018). Leaf mine. On Dichanthelium, eggs are inserted near the leaf margin, associated with a cluster of feeding punctures. The usually gregarious larvae initially mine along the margin toward the apex, then turn around and form a large, dirty whitish upper surface blotch with generally distributed, liquidy, greenish to brown frass. Concentric feeding lines are present in some mines. Mines on Glyceria may be on the lower leaf surface and the photographed examples do not have associated feeding punctures. Puparium. Yellowish-brown to reddish-brown; formed outside the mine. Phenology and voltinism. In Massachusetts, larvae feeding from late June to mid-July emerge as adults from mid-July to early August; larvae collected in mid-August yielded some adults in early September, with some females emerging the following spring (Eiseman & Lonsdale 2018). In Iowa, a larva found feeding in late July emerged as an adult about three weeks later. In North Carolina, larvae collected in early May did not emerge as adults until the following spring, whereas a larva collected in early June yielded an adult later that month. Distribution. USA: *IA, MA, NC, TN., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 6, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Melanagromyza rudbeckiae Eiseman & Lonsdale, spec. nov. (Figs. 13–15, 105–110) Holotype. USA. IOWA: Winneshiek Co., Meadow Farm, 20.viii.2017, em.?, J. van der Linden, ex Rudbeckia laciniata, # CSE4664, CNC1135655 (1♁). Paratypes. USA. WISCONSIN: Grant Co., Thomas Wet. Prairie, 23.ix.1997, A.H. Williams, stems of Rudbeckia lacinata, T7 N R1 W Sect. 7, [host stem] stripped of leaves and inflorescences, put into sterile containers over sterile soil and netted w/hosiery, outdoors until 2.iii.1998 when tightly caged in lab, em. 15–24.iv.1998, CNC934510–934512 (2♁, WIRC; 1♁, CNC). Etymology. The specific epithet refers to the host plant genus, Rudbeckia L. Host. Asteraceae: Rudbeckia laciniata L. Larval biology. Internal stem borer. Puparium. Formed within the stem. Phenology and voltinism. Pupae overwinter, with adults emerging after six to eight weeks of exposure to warm temperatures in spring. The holotype was reared from a larva or pupa collected in mid-August, but unfortunately its emergence date was not recorded. Distribution. USA: IA, WI. Adult description. Wing length approximately 2.3–2.5 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 0.7–0.8. Eye height divided by gena height: 3.6–4.8. First flagellomere small, rounded. Orbital plate slightly projecting. Gena horizontal on posterior half, angled upwards on anterior half. Face with very shallow ridge. Clypeus rounded. Head distinctly higher than long, not as developed anteriorly as in most congeners; many fibers trapped under setae on head, obscuring much of frons. Thorax subshining. Right foreleg and left hindleg missing. Chaetotaxy: Three ori (anterior seta nearly flat on frons), two ors. Orbital setulae in several rows, mostly erect to proclinate on outer row, mostly reclinate on inner. Eye setose dorsomedially, relatively dense in paratypes, sparse in holotype. Postvertical seta slightly longer than ocellar. Two strong dorsocentral setae. Acrostichal setulae in eight rows. One smaller additional seta on katepisternum. Mid tibia with two posteromedial setae, only one on left leg of holotype. Coloration: (Figs. 13–15) Setae dark brown. Color dark brown, including halter; slight metallic green shine on dorsum of thorax that is also evident and slightly more pronounced on abdomen. Calypter margin and hairs yellow. Genitalia: (Figs. 105–110) Epandrium with minute posteroventral spine. Surstylus shallow, wide, curved, fused to anteroventral surface of epandrium; inner distal margin with scattered tubercle-like setulae. Cercus large, welldeveloped. Hypandrium relatively wide, sides slightly converging apically to narrower apical apodeme; inner lobe Y-shaped with sockets on anterior process. Phallophorus with base abruptly constricted, short and with sides flared; swollen distoventrally. Basiphallus U-shaped, long dorsally, pointed at ends; distant from basiphallus+mesophallus, with intervening gap nearly as long as phallophorus. Mesophallus short, cylindrical, tapering to ventromedial point of fusion on distiphallus; base slightly exceeding that of distiphallus. Distiphallus relatively short compared to congeners, subovate in ventral view with length twice width; one pair of ventrolateral tubules flanking mesophallus; dark, thick ventromedial plate greatly swollen immediately past mesophallus; dorsal chamber small, short, widest apically with margin flared laterally, internally with one pair of spinulose pads; tubular process arising from dorsal chamber short and straight. Ejaculatory apodeme with narrow, pale blade with medial rib, short stem with long lateromedial process, and long weak tube arising from base on side opposite duct; sperm pump with wide basal sclerotization, including transverse bar that is upcurved at ends. Comments. Another Nearctic species, Melanagromyza minimoides Spencer, has been reared from Rudbeckia laciniata, but the larvae feed in the seedhead rather than boring in the stem (Spencer & Steyskal 1986; this paper), and the calypter hairs are dark brown, not yellow. Melanagromyza rudbeckiae is the undetermined species Williams (1999) reported rearing from overwintering stems of R. laciniata in Wisconsin, and his specimens are included here as paratypes. Melanagromyza rudbeckiae is a relatively small, faintly greenish species with a characteristic phallus that has the distiphallus and basiphallus widely separated, the mesophallus base slightly exceeds that of the distiphallus, and the distiphallus is relatively short with the medial tubule well exposed apically, the venter is thickly sclerotized, and in ventral view the outline is narrowly egg-shaped with the sides slightly flared distolaterally. Unlike similar Melanagromyza identifiable from couplet 36 in Spencer & Steyskal (1986), M. rudbeckiae has a relatively long space between the basiphallus and the distiphallus. Melanagromyza longensis is slightly larger (wing length 2.6 mm) but otherwise externally similar (including three ori), but the distiphallus is approximate to the basiphallus. Other similar but slightly larger Nearctic species are either bluish in color (M. hicksi Steyskal) or have a projecting frons (M. angelicae)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 11-12, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","Williams, A. H. (1999) Fauna overwintering in or on stems of Wisconsin prairie forbs. Proceedings of the North American Prairie Conference, 16, 156 - 161."]}

Haplopeodes loprestii Eiseman & Lonsdale, spec. nov. (Figs. 32–39, 133–139) Holotype. USA. CALIFORNIA: San Diego Co., Coyote Canyon, 14.iii.2018, em. 18.v–3.vi.2018, E. LoPresti, ex Fagonia laevis, # CSE4545, CNC1135597 (1♁). Paratype. Same data as holotype, CNC1135598 (1♀ ) . Etymology. This species is named for Eric F. LoPresti, ecologist and evolutionary biologist, who collected the host material from which it was reared. Host. Zygophyllaceae: Fagonia laevis Standl. Larval biology. Apparently a leafminer. The host material was collected to rear leaf-mining larvae of a gelechiid moth, and the presence of the flies only became evident when the puparia appeared. The rearing vial also contained fruits and stems, so the possibility of these as feeding sites cannot be entirely excluded, but as far as is known all Haplopeodes species are leafminers. Puparium. (Fig. 34) Pale, straw-colored; formed externally. Phenology and voltinism. The two known specimens were reared from larvae that fed in March, pupating by the 20 th and emerging as adults four to six weeks later. Distribution. USA: CA. Adult description. Wing length 1.2 mm (♁), 1.4 mm (♀). Vein dm-m absent. Eye height divided by gena height: 3.2–3.5. First flagellomere slightly longer than high, apically rounded. Arista short, only 1.5 times length of first flagellomere. Orbital plate shallow but distinctly projecting when viewed laterally, especially anteriorly on head; continuing as modestly developed ring around eye. Seen dorsally, frons wider than eye, slightly broader anteriorly. Anterior ocellus slightly displaced anteriorly. Epistoma large, broad, subrectangular, resulting in correspondingly shorter face; face with shallow medial carina. Palpus small, narrow, length approximately three times width. Thorax subshining. Wing veins costalized (closely spaced anteriorly), with width of cell r 4+5 subequal to that of cell sc; length of costal cell subequal to length of cell r 1 past insertion of vein R 1. Chaetotaxy: Setae short. Three ori, very gracile; ors absent. Few orbital setulae, slightly inclinate and erect, partially proclinate. Ocellar seta straight, approximately ¾ length of ori. Postvertical seta absent. Five dorsocentral setae, strongly decreasing in length anteriorly; female with additional small setula in dorsocentral row anteriorly. Only one notopleural seta, medially positioned. Notal setulae mostly absent; female with 1 anteromedial acrostichal setula and with few setulae in intra-alar region. One pair of lateral scutellar setae and two pairs of apical scutellar setae (left anterior seta absent in male). Katepisternal seta not visible in male. Coloration: (Figs. 32–33, 35–39) Setae brown to dark brown with pale brown shine. Base color light yellow with head paler. First flagellomere brown with inner surface (not including margins) yellow; ocellar triangle dark brown; vertex light yellow; posterolateral corner of frons with brown patch fading to base of inner vertical seta; face deeper yellow with slight orange tint; venter of gena with brownish line; back of head dark brown with dorsomedial region paler and venter yellow. Notum dark brown to brown; center of scutellum and mediotergite slightly paler brown; postpronotum light brown, becoming yellower anteriorly and laterally; notopleuron and supra-alar spot light brown to yellow, with notopleuron yellower medially; katatergite light brown. Proepisternum light brown; anepisternum with anteroventral corner paler yellow, bordered posteriorly by oblique brown stripe; katepisternum with brown spot below level of seta; anepimeron irregularly brown; meron mostly dark brown. Legs light yellow; base of coxae narrowly brown; femora with indistinct brownish tint on apical ¼–½; tibiae brown with mediolateral and medioventral regions broadly yellow; tarsi brown. Abdomen brown with lateral margin of tergites narrowly yellow; epandrium darker brown. Calypter margin and hairs yellow. Variation. Female: Darker than male. Setae darker brown to blackish. Only inner-basal surface of first flagellomere yellow; scape and pedicel yellow. Posterolateral corner of frons darker. Paler parts of scutum brown tinted. Anepisternum, anepimeron and meron brown; base color of katepisternum brownish-orange. Femora extensively brown mottled. Abdominal tergites entirely brown. Genitalia: (Figs. 133–139) Epandrium shallow. Surstylus short, incurved, apical margin irregular but essentially straight, without tubercle-like setae or narrow protrusions referred to as “teeth” in Steyskal (1980a); base (concealed within epandrium) incurved. Cerci weakly sclerotized, much thicker laterally, closely spaced. Hypandrium with short, semicircular arch; inner lobe weakly sclerotized, more so on distal margin bearing single seta; arm long, well-defined. Postgonite dark, narrow, with single subapical seta. Phallus small and weakly sclerotized, but comparatively large and dark for Haplopeodes; phallophorus cylindrical, fused to curved basiphallus, which is represented by short ventral sclerotization that is slightly elongated on left side. Ejaculatory duct strongly upcurved past basiphallus, with exposed portion approximately as long as basiphallus; apex rounded, dorsoventrally flattened and lightly sclerotized. Ejaculatory apodeme narrow but well-developed, darker stem grading into blade. Comments. This is the first record of any agromyzid from Fagonia, and the only agromyzids previously recorded from Zygophyllaceae are extremely polyphagous Liriomyza spp. Known hosts of Haplopeodes are otherwise restricted to Amaranthaceae, Portulacaceae, and Solanaceae (Benavent-Corai et al. 2005). While minor color differences between the sexes are evident in some Haplopeodes, the sexual dimorphism observed here is pronounced and unique, with the female being much darker than the male. Both sexes are considerably darker than other Haplopeodes, with the scutellum entirely dark brown (unique among Nearctic Haplopeodes), the first flagellomere and scutellum brown, the pale grey pruinosity usually observed on the thorax is absent (the scutum is subshining), the tibiae are brown with yellow medial regions and the femora are brown mottled. Furthermore, the acrostichal setulae are virtually absent, the orbital plate and parafacial are produced, an epistoma is present, the radial veins are crowded anteriorly, the phallus is pigmented apically and surstylus spines are absent., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 22-23, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Steyskal, G. C. (1980 a) Haplopeodes, a new genus for Haplomyza of authors (Diptera, Agromyzidae). Proceedings of the Entomological Society of Washington, 82 (1), 140 - 151.","Benavent-Corai, J., Martinez, M. & Jimenez Peydro, R. (2005) Catalogue of the host plants of the world Agromyzidae (Diptera). Bollettino di Zoologia agraria e di Bachicoltura, Series II, 37 (Supplementum), 1 - 97."]}

Melanagromyza hieracii Eiseman & Lonsdale, spec. nov. (Figs. 9–12, 61, 99–104) Holotype. USA. IOWA: Allamakee Co., Fish Farm Mounds WMA, 30.ix.2017, em. spring 2018, J. van der Linden, ex Hieracium scabrum, # CSE4669, CNC1144040 (1♁). Etymology. The specific epithet refers to the host plant genus, Hieracium L. Host. Asteraceae: Hieracium scabrum Michx. Larval biology. (Fig. 61) The larva bores in the pith of the stem, forming a linear gallery. Puparium. (Fig. 12) Straw-colored, formed at the end of the larval gallery; rear spiracular horns pronounced, blackish, curved and pointed. Phenology and voltinism. The holotype emerged after overwintering from a puparium collected on 30 September. All other puparia found at that time in stems of H. scabrum were already empty. Assuming these represented the same species, this suggests either that Melanagromyza hieracii has more than one generation or that adults may be capable of overwintering. Distribution. USA: IA. Adult description. Wing length 2.7 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 0.7. Eye height divided by gena height: 4.5. First flagellomere small, rounded. Ocellar triangle largely obscured by detritus, appearing to reach past posterior ori. Anterior half of gena slightly angled upwards. Cheek not evident. Clypeus rounded. Eye and head longest above midpoint. Thorax subshining. Body in relatively poor condition; partially collapsed, some setae missing. Chaetotaxy: Four gracile ori strongly incurved (nearly flat on frons), two strong ors; setae long. Several rows of scattered orbital setulae, as long as eye hairs, erect (inner setulae) to reclinate (outer). Ocellar setae possibly missing, bases obscured. Eye hairs sparse and scattered on dorsal half with dense dorsomedial patch. Anterior of two dorsocentral setae missing, sockets large. Acrostichal setulae in ten irregular rows. Katepisternum with one additional subdominant seta. Mid tibia with two posteromedial setae. Coloration: (Figs. 9–11) Setae dark brown. Body dark brown, including halter; notum faintly green metallic; abdomen green metallic with blue tint; wing veins brown. Calypter margin and hairs yellow. Genitalia: (Figs. 99–104) Epandrium with posteroventral spine. Surstylus fused to anteroventral margin of epandrium, relatively narrow and produced with slight posterodistal extension; distal margin with irregular rows of tubercle-like setae. Cercus large and well-developed. Hypandrium broadly rounded basally, with long narrow apical apodeme; inner lobe U-shaped with minute setulae. Phallophorus swollen ventrally, base much narrowed. Basiphallus U-shaped (right side irregularly sclerotized in dissected male), with short gap between apex and mesophallus. Mesophallus swollen cylindrical, narrowest at point of insertion to distiphallus, which is relatively posterobasal. Distiphallus short, globular, with short ventromedial swelling into which mesophallus inserts; basal section mostly made up of posteriorly convergent tubules flanking mesophallus; ventral plate subquadrate, distal margin slightly flared laterally; dorsal chamber shifted distally, widest subapically, with one pair of minutely spinulose internal pads; tubular process narrow, distal half exposed, sinuate when viewed laterally. Ejaculatory apodeme with short dark stem with long lateromedial process; blade narrow with medial rib, margin along one side narrow and irregular (broken?), other side wider and rounded with margin clear; sperm pump with dark transverse bar upcurved at ends. Comments. This is the first record of a North American Melanagromyza from Hieracium. The European M. oligophaga Spencer includes H. umbellatum L. among its hosts (Spencer 1990). Externally, Melanagromyza hieracii is not unlike many other congeners with a metallic green shine (weak on the thorax and with a blue tint on the abdomen), a white calypter, two dorsocentrals and a shallow orbital plate. The key in Spencer & Steyskal (1986) will bring the species to M. longensis Spencer / M. virginiensis Spencer, but unlike these species, it has four ori. The phallus is unlike those of these two species in that the dorsal chamber of the distiphallus is parallel-sided, the lateral margins of the distiphallus are very strongly flared apically, the medial tubule is pronounced and long, far exceeding the apex of the short dorsal chamber, and the mesophallus is strongly shifted basally on the distiphallus. The combination of these phallic features, especially the flared distolateral margins of the distiphallus and the basally shifted mesophallus, is quite unusual for the genus. Also see comments for Melanagromyza urticae., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 9-10, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. (1990) Host specialization in the world Agromyzidae (Diptera). Series entomologica, 45, i-xii + 1 - 444. https: // doi. org / 10.1007 / 978 - 94 - 009 - 1874 - 0","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Liriomyza trifolii (Burgess) (Fig. 80) Material examined. NORTH CAROLINA: Scotland Co., Laurinburg, St. Andrews University, 20.v.2018, em. 4.vi.2018, T. S. Feldman, ex Mecardonia acuminata, # CSE4579, CNC1135666–1135670, CNC1144089 (3♁ 3♀). Hosts. (Note: underlined hosts are from a greenhouse experiment.) Amaranthaceae: Amaranthus palmeri S.Watson, Beta vulgaris L., Chenopodium album L., Spinacia oleracea L.; Amaryllidaceae: Allium cepa L.; Apiaceae: Apium graveolens L., Daucus carota L.; Araliaceae: Hydrocotyle umbellata L., H. verticillata Thunb.; Asteraceae: Ageratum L. (Heinz & Parrella 1990), [Ambrosia artemisiifolia L.], “ Aster ” (cultivated), Baccharis halimifolia L., Bidens alba (L.) DC., B. pilosa L., Callistephus chinensis (L.) Nees, Chrysanthemum × morifolium (Ramat.) Hemsl., Conoclinium coelestinum DC., [Cosmos bipinnatus Cav.], Dahlia Cav., Erechtites hieraciifolius (L.) Raf. ex DC., Eupatorium capillifolium (Lam.) Small ex Porter & Britton, E. serotinum Michx., Flaveria trinervia (Spreng.) C.Mohr, Gaillardia aristata Pursh, Galinsoga quadriradiata Ruiz & Pav., Gamochaeta pensylvanica (Willd.) Cabrera, Gerbera jamesonii Bolus, Helianthus annuus L., Hymenopappus scabiosaeus L’Hér., Lactuca canadensis L., L. sativa L., [Leucanthemum vulgare Lam.], Melanthera nivea (L.) Small, Mikania scandens (L.) Willd., Packera glabella (Poir.) C.Jeffrey, Parthenium hysterophorus L., “ Senecio ”, Sonchus asper (L.) Hill, S. oleraceus L., Symphyotrichum cordifolium (L.) G.L.Nesom, Synedrella nodiflora (L.) Gaertn., Tagetes erecta L., Taraxacum officinale F.H.Wigg., Tridax procumbens L., [Vernonia gigantea (Walter) Trel.], Xanthium L., Zinnia L.; Caryophyllaceae: Gypsophila L.; Cucurbitaceae: Cucumis melo L., C. sativus L., Cucurbita pepo L., Melothria pendula L.; Fabaceae: Crotalaria incana L., [Lathyrus japonicus Willd.], Medicago lupulina L., M. sativa L., Phaseolus lunatus L., P. vulgaris L., Pisum sativum L., Trifolium repens L., Vicia sativa L., Vigna luteola (Jacq.) Benth., V. radiata (L.) R.Wilczek, V. unguiculata (L.) Walp.; [Lamiaceae: Glechoma hederacea L.]; Malvaceae: Abelmoschus esculentus (L.) Moench, “ Hibiscus ”, Malva moschata L.; Nyctaginaceae: Abronia villosa S.Watson; Plantaginaceae: * Mecardonia acuminata (Walter) Small, Plantago major L.; [Poaceae: Avena sativa L.; Polygonaceae: Fallopia convolvulus (L.) Á.L̂ve, Persicaria maculosa Gray.]; Ranunculaceae: Ranunculus repens L.; Solanaceae: Capsicum annuum L., Petunia Juss., Physalis angulata L., P. philadelphica Lam. (Pérez-Alquicira et al. 2019), P. pubescens L., Solanum americanum Mill., S. dulcamara L., S. lycopersicum L., S. melongena L., S. nigrum L., S. tuberosum L.; [Turneraceae: Piriqueta cistoides subsp. caroliniana (Walter) Arbo]; Verbenaceae: Verbena L.; Zygophyllaceae: Kallstroemia maxima (L.) Hook. & Arn., Tribulus terrestris L. (Eiseman & Lonsdale 2018 and references therein; see Lonsdale (2011) for additional host genera recorded outside of North America). Leaf mine. (Fig. 80) Narrow and linear throughout; greenish to whitish, with dark green to black frass, typically in alternating strips or closely spaced grains, but in some cases partially forming a squiggly line. Mines are generally confined to the upper leaf surface, but some mines on Mecardonia were partly formed on the lower surface. Puparium. Yellowish; formed outside the mine. Phenology and voltinism. This species is multivoltine. We have collected larvae on 7 March in southern California, 26 March in Florida, and 20 May and 2 June in North Carolina, with adults emerging in 12–25 days in each case. Distribution. Widespread in North, Central, and South America; introduced in the Old World. Largely restrict- ed to greenhouses in colder temperate regions (Lonsdale 2011). Comments. This is the first record of any agromyzid from Mecardonia Ruiz & Pav., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 32, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Heinz, K. M. & Parrella, M. P. (1990) Holarctic distribution of the leafminer parasitoid Diglyphus begini (Hymenoptera: Eulophidae) and notes on its life history attacking Liriomyza trifolii (Diptera: Agromyzidae) in chrysanthemum. Annals of the Entomological Society of America, 83 (5), 916 - 924. https: // doi. org / 10.1093 / aesa / 83.5.916","Perez-Alquicira, J., van der Knaap, E., Mercer, K., McHale, L., Mitchell, T., Luna-Ruiz, J., Texocotitla-Vazquez, E., Vargas- Ponce, O. & Michel, A. (2019) Genetic structure of Liriomyza trifolii (Diptera: Agromyzidae) associated with host plants from southeastern Mexico. Environmental Entomology, 48 (1), 253 - 262. https: // doi. org / 10.1093 / ee / nvy 184","Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Lonsdale, O. (2011) The Liriomyza (Agromyzidae: Schizophora: Diptera) of California. Zootaxa, 2850 (1), 1 - 123. https: // doi. org / 10.11646 / zootaxa. 2850.1.1"]}

Melanagromyza sp. 2 Material examined. OKLAHOMA: Payne Co., Mehan, 36.013839°, -96.998103°, 12.vii.2017, em. ~ 10.iv.2018, M.W. Palmer, ex Lactuca canadensis, # CSE4452, CNC1135642 (1♀). Host. Asteraceae: Lactuca canadensis L. Larval biology. Probably an internal stem borer (see Comments). Puparium. Formed within the stem. Phenology and voltinism. This species is evidently univoltine, given that the adult emerged in spring from a stem collected the previous July. Comments. The only previous record of a Melanagromyza from Lactuca is that of several M. splendida Frick paratypes from Hawaii, listed as “reared from larvae mining lettuce leaves” (Frick 1953)—which should not be construed as representing true leafmining since as far as is known M. splendida feeds as a borer in the stem or in the midrib of large leaves (Spencer 1973). Unlike M. splendida, the Oklahoma female has closely spaced fronto-orbitals (the anterior pair is not anteriorly displaced) and the wing is approximately 3.2 mm (not 1.9–2.6 mm). The female emerged from a bulk sample of mined stems, with the mines terminating in conspicuous gall-like “scabs” of stem tissue and dried latex, inside which the puparia were formed. We are reasonably sure that these external symptoms were in fact caused by an undetermined Ophiomyia species, and we presume that the Melanagromyza larva fed and pupated in the pith of a stem., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 15, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Frick, K. E. (1953) Further studies on Hawaiian Agromyzidae (Diptera) with descriptions of four new species. Proceedings of the Hawaiian Entomological Society, 15 (1), 207 - 215."]}

Melanagromyza verbenivora Eiseman & Lonsdale, spec. nov. (Figs. 20–23, 66–67, 117–121) Holotype. USA. IOWA: Winneshiek Co., Beard Farm, 12.vii.2017, em. summer 2017, J. van der Linden, ex Verbena stricta, # CSE4648, CNC1144043 (1♁). Paratypes. IOWA: same data as holotype, # CSE4649, CNC1135656 (1♀); same data as holotype, em. 14.vii.2017, # CSE4653, CNC1144047 (1♁ [illustrated]); Winneshiek Co., Beard Farm, vi–vii.2017, em. summer 2017, J. van der Linden, ex Verbena stricta, # CSE4657, CNC1144042 (1♀); Decorah, Trout Run Trail, 43°18’5.04”N 91°48’6.60”W, 12–22.vii.2017, em. late vii—early viii.2017, J. van der Linden, ex Verbena stricta, # CSE4942, CNC1643670–1643672 (1♁ 2♀). Etymology. The specific epithet refers to the host plant genus, Verbena L. Host. Verbenaceae: Verbena stricta Vent. Larval biology. (Figs. 66–67) Some larvae bore in the stem, forming a spiral gallery that is partly visible externally. This causes some stunting of the growing shoot and wilting of the terminal portion (Fig. 66). Others bore in the rachis of more developed shoots, resulting in a somewhat thickened seedhead with a shriveled, blackened tip (Fig. 67). The interior of the stem or rachis is filled with granular frass. Puparium. (Fig. 23) Whitish; formed within the larval feeding site. Phenology and voltinism. Larvae feed in early summer, with adults emerging beginning in mid-July. Later generations have not been observed. Distribution. USA: IA. Adult description. Wing length 2.4–2.6 mm (♁), 2.7–2.9 mm (♀). Length of ultimate section of vein M 4 divided by penultimate section: 0.7. Eye height divided by gena height: 4.8–5.3. First flagellomere small, rounded. Lunule extending to level of middle ori. Ocellar triangle slightly shinier than vitta, extending to level of middle or posterior ori. Facial carina shallow, sharp. Gena shallowly upcurved anteriorly. Cheek narrow, only evident on anterior half. Clypeus rounded. Head (seen laterally) longest at midpoint. Eye tapering to a point posterodorsally. Thorax subshining. Chaetotaxy: Three ori, two ors; spacing subequal with anterior ori sometimes slightly distant; setae becoming slightly more slender and short anteriorly. Ocellar and postvertical setae subequal to posterior ori. Eye sparsely short setose on dorsal half; female with hairs slightly denser dorsally, male with longer, denser patch of hairs dorsomedially. Orbital setulae in several scattered rows; erect to slightly reclinate. Two strong dorsocentral setae. Acrostichal setulae in ten irregular rows. Katepisternum with two additional smaller, closely spaced setae dorsomedially. Mid tibia with two posteromedial setae. Coloration: (Figs. 20–22) Setae dark brown. Body dark brown, including halter; thorax and abdomen metallic green with blue tint that is stronger on abdomen; blue color dominant in one female. Calypter margin and hairs yellow. Genitalia: (Figs. 117–121) Epandrium with small posteroventral spine. Surstylus fused to anteroventral margin of epandrium, very short and wide with 1–3 irregular rows of tubercle-like setae along inner surface. Cercus large, well-developed. Hypandrium subtriangular with apex slightly narrowed; inner lobe U-shaped and irregular with several sockets. Phallophorus (damaged in dissected male) tapering basally, venter bulging. Basiphallus U-shaped, nearly reaching mesophallus. Mesophallus cylindrical, slightly curved, narrowed to ventromedial point of fusion on distiphallus; base of mesophallus and distiphallus level. Distiphallus pear-shaped in ventral view, with narrow ventromedial plate swollen immediately past mesophallus; ventrolateral tubules narrow, largely hidden behind mesophallus along midline; dorsal chamber widest dorsally, broadly opened apically, with one pair of thick spinulose dorsal pads on inner surface; tubular inner process relatively wide, not far exceeding ventral plate. Ejaculatory apodeme not found. Comments. The only previous record of a North American Melanagromyza from Verbena was that of an unemerged male of an apparently undescribed species that was said to be entirely black, extracted from a stem of V. scabra Vahl in Florida (Spencer & Stegmaier 1973). It was noted that “The posterior spiracles of the puparium each bear ten bulbs, without a central horn; the two processes are separated by twice their own diameter.” Unfortunately no puparia of M. verbenivora have been preserved for comparison. Melanagromyza verbenae Spencer was described from adults caught on V. litoralis Kunth in Chile, and due to the general similarity of the male genitalia with those of the Florida specimen, Spencer (1982) considered it certain that this plant was the larval host and presumed that the Chilean species is likewise an internal stem borer. Spencer (1990) referred to the two species as “related or possibly identical”. Using Spencer & Steyskal (1986), some specimens of Melanagromyza verbenivora key to M. longensis, which also has three ori and a similar phallus, but in that species, the arista appears bare (not minutely pubescent), there are about eight rows of acrostichal setulae, the basiphallus nearly forms a complete ring, and the dorsal chamber of the distiphallus is more rounded dorsally (not flat) and constricted apically (not broadly open). Other specimens will key to M. verbesinae, but in this species there are two ori, the ocellar triangle does not reach the posterior ori, and the distiphallus is widest apically. The gena is slightly angled forward as in M. buccalis Spencer, but not as prominent; M. buccalis is slightly smaller (wing length 1.9–2.5mm), there are two to four ori (widely spaced if only two), there are only eight rows of acrostichal setulae and the phallus is narrower., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 13-14, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Stegmaier, C. E. Jr. (1973) Agromyzidae of Florida with a supplement on species from the Caribbean. Arthropods of Florida and Neighboring Land Areas, 7, 1 - 205. https: // doi. org / 10.1007 / 978 - 94 - 017 - 0683 - 4 _ 1","Spencer, K. A. (1982) Agromyzidae (Diptera) in Chile. Stuttgarter Beitrage zur Naturkunde, Serie A (Biologie), 357, 1 - 55.","Spencer, K. A. (1990) Host specialization in the world Agromyzidae (Diptera). Series entomologica, 45, i-xii + 1 - 444. https: // doi. org / 10.1007 / 978 - 94 - 009 - 1874 - 0","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Phytomyza flavilonicera Eiseman & Lonsdale, spec. nov. (Figs. 48–52, 82–83, 148–153) Holotype. USA. OKLAHOMA: Payne Co., Stillwater, Bustani Plant Farm, 22.iv.2018, em. 4–14.v.2018, M.W. Palmer, ex Lonicera sempervirens, # CSE4480, CNC1144151 (1♁). Paratypes. OKLAHOMA: same data as holotype, CNC1144152–1144157 (3♁ 3♀); Payne Co., Stillwater, Bustani Plant Farm, 22.iv.2018, em. 29.iv.2018, M.W. Palmer, ex Lonicera flava, # CSE4614, CNC1135622–1135624 (3♀); same data, # CSE4621, CNC1135639–1135640 (2♀); em. 5.v.2018, # CSE4623, CNC1144147–1144150 (1♁ 3♀); Bustani Plant Farm, 22.iv.2018, em. 22–25.iv.2018, M.W. Palmer, ex Lonicera sempervirens, # CSE4450, CNC1144144, CNC1144146 (1♁ 1♀); em. 29.iv.2018, # CSE4616, CNC1135618–1135621 (1♁ 3♀); em. 6.v.2018, # CSE4618, CNC1135610–1135618 (5♁ 4♀); Sanborn Lake, 9.vi.2017, em. 18.vi.2017, M.W. Palmer, ex Lonicera sempervirens, # CSE3940, CNC939928 (1♀). Etymology. The specific epithet is a noun in apposition, a portmanteau referring to the yellow head of the adult (L. flavus —yellow) and the host plant genus, Lonicera L. Hosts. Caprifoliaceae: Lonicera flava Sims, L. sempervirens L. Leaf mine. (Figs. 82–83) Whitish, on the upper leaf surface; initially stellate, and in some cases remaining a small, digitate blotch throughout development; in other cases one or a few linear branches emanate from the stellate beginning. Frass forms a dark central patch in the stellate portion and is otherwise deposited in closely spaced grains or beaded strips. Puparium. (Fig. 52) Whitish to pale brown, with a broad, dark, elongate central band on the ventral surface; formed within the mine, with the ventral surface against the upper epidermis. Phenology and voltinism. This species is probably multivoltine. Leaf mines have been collected in April with adults emerging from late April to mid-May, as well as in early June with an adult emerging in mid-June. Distribution. USA: OK. Adult description. Wing length 1.5–1.6 mm (♁), 1.7–1.8 mm (♀). Vein dm-m absent. Eye height divided by gena height: 3.7–4.2. First flagellomere slightly longer than high, stout and appearing somewhat enlarged with dorsal and ventral margins parallel for short distance medially. Orbital plate projecting along anterodorsal eye margin (seen laterally). Cheek distinct. Posterior ocelli slightly displaced. Thorax pruinose (see below). Chaetotaxy: Two ors, two ori, decreasing in length anteriorly; anterior ori sometimes setula-like; sometimes smaller third ori present on one side posteriorly; one ors sometimes missing. Postvertical seta subequal to posterior ors. Ocellar seta slightly shorter than postvertical. Four dorsocentral setae, decreasing in length anteriorly. Approximately six scattered rows of acrostichal setulae. Setulae on pleuron nearly absent. Coloration: (Figs. 48–51) Setae dark brown. Head mostly light yellow; first flagellomere dark brown, scape and pedicel deep yellow; back of head, vertex (slightly paler), broad spot around ocellar tubercle and posterolateral corner of frons past base of inner vertical seta dark brown; orbital plate dark brown posteriorly with stripe extending to surround base of posterior ors and paler stripe with slight silvery pruinosity extending to meet base of anterior ors; slight mottling at base of posterior ori; remainder of orbital plate and frontal vitta dirty yellow; clypeus greyish-brown, very narrow and pale medially; palpus brown. Thorax dark brown dorsally, paler on pleuron; with bluish-grey pruinosity that is dense on notum and with a coppery tint posteriorly on scutum and scutellum; yellow mottling along notopleural sutures and postalar wall; anterior spiracle and vertical line dorsal to it yellow; anepisternum sometimes with margins irregularly yellow; yellow above coxopleural streak; halter yellow. Wing veins light brownish-yellow. Legs brown; apices of fore and mid coxae yellow; apices of femora yellow for distance equal to width of femur apex; bases of tibiae yellow (spot narrower on posterior legs) and dorsal apex of fore tibia yellow; fore tarsus brownish-yellow. Abdomen brown with lateral margin of tergites yellow; epandrium paler brown with anterior and posterior margins yellow. Calypter yellow with hairs light brown to golden. Variation. Female: Pigmented line along vertex paler, usually at least at base of postvertical setae. Apex of fore tibia faintly yellow; fore tarsus light brown. Abdominal tergites entirely brown or with very narrow yellow margin restricted to tergites 1–3. Calypter hairs light brown to brown. Genitalia: (Figs. 148–153) Surstylus small, rounded, incurved, short setose; fused to epandrium. Hypandrium subtriangular with broad, rounded apex; inner lobe narrow, V-shaped with two apical setae. Postgonite narrow with one seta. Phallophorus simple, cylindrical, with one pair of narrow band-like lateral lobes. Basiphallus consisting of one pair of long narrow plates with irregular margins and tapered apices; right plate with base positioned dorsally, left plate with base wrapping around venter. Hypophallus (one pair of lateral sclerites) small, irregular in outline, longer than wide; some dissected males with sclerite atrophied to entirely absent. Mesophallus not evident. Distiphallus short, sharply angled dorsally, narrow; when viewed ventrally, with wider, rounded base that splits into one pair of narrow subparallel bands that slightly diverge apically and appear slightly bifid with outer arm weaker and shorter. Ejaculatory apodeme very small, weakly sclerotized, with long weak tubule emerging from base opposite duct; sperm pump clear. Comments. The leaf mine of Phytomyza flavilonicera is similar to that of P. sempervirentis Eiseman & Lonsdale, which also occurs on Lonicera sempervirens, but the latter species usually moves to the lower leaf surface before pupating, and the dark ventral area on its puparium is usually a narrow stripe rather than a broad band (Eiseman & Lonsdale 2018). However, a pair of mines photographed in Missouri (Hartley 2017) seems to be within the range of variation of both species. The two are not known to be sympatric, with P. flavilonicera found only in Oklahoma and P. sempervirentis recorded from Oregon, Massachusetts, North Carolina, and Alabama, but more collecting of these mines from Midwestern states will be needed to clarify the extents of their distributions. Twelve parasitoids reared from Phytomyza flavilonicera were identified by J.T. Huber as Diglyphus websteri (Crawford) (Eulophidae), a specialist on Agromyzidae but with a wide host range within this family. Phytomyza flavilonicera is a relatively pale Caprifoliaceae-mining Phytomyza, being yellowish laterally on the scutum (the central disc is bluish-grey pruinose), and paler on the pleuron and legs with the knees yellowish. The head is also mostly pale, having a yellow face, a dirty yellow frontal vitta and anteriorly yellowish orbital plate. It is further characterized by having numerous rows of acrostichals, a small to absent subovate sclerite on the hypophallus, and a relatively small, simple, upturned distiphallus. In ventral view, the outline of the distiphallus differs from that of similar species such as P. nigrilineata in being very narrow with a broader lobate base, and it is medially split past the insertion point of the duct, with the points slightly diverging, fading and appearing partially bifurcated apically. This distiphallus structure is similar to some P. sempervirentis Eiseman & Lonsdale (see Eiseman & Lonsdale 2018: Figs. 358, 359), but this is a darker species with a brown head, the distiphallus is much more wedge-shaped in appearance laterally and in ventral view a dark V-shape predominates., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 34-36, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Hartley, C. (2017) Leaf miner on Passiflora. Available from: https: // bugguide. net / node / view / 1386669 (accessed 14 December 2019)"]}

Phytomyza triostevena Eiseman & Lonsdale, spec. nov. (Figs. 53–56, 86, 154–158) Holotype. USA. IOWA: Allamakee Co., 5.ix.2017, puparium removed from midrib 26.ix.2017, adult em. by spring 2018, J. van der Linden, ex Triosteum, # CSE4663, CNC1144064 (1♁). Etymology. The specific epithet is a noun in apposition, a portmanteau referring to the position of the larval mines in the veins and midribs of leaves of Triosteum L. (L. vena —a vein). Host. Caprifoliaceae: Triosteum L.; similar leaf mines have been found on T. aurantiacum E.P.Bicknell (Mews 2019a). Leaf mine. (Fig. 86) The larva initially forms a narrow, serpentine mine in the leaf blade. It soon enters a lateral vein and follows this to the midrib, in which it feeds until mature. Puparium. (Fig. 56) Brown; formed within the midrib, near the base of the leaf. Phenology and voltinism. Leaf mines have been observed beginning in early September, with adults probably not emerging until the following spring. Distribution. USA: IA; similar leaf mines have been found in Canada: ON (Mews 2019a). Adult description. Wing length 1.8 mm (♁). Female unknown. Vein dm-m absent. Eye height divided by gena height: 3.5. First flagellomere rounded, as long as high. Orbital plate well-defined, pruinose, narrowing anteriorly. Frontal vitta minutely pitted, appearing darker. Cheek well-developed. Thorax subshining. Chaetotaxy: Two ori (anterior seta slightly shorter and positioned closer to eye margin); one ors. Ocellar seta subequal to anterior ori; postvertical subequal to ors. Four dorsocentral setae, decreasing in length anteriorly, with anterior two much shorter (less than half length of first seta). Approximately four to five irregular rows of acrostichal setulae. Coloration: (Figs. 53–55) Setae dark brown. Body dark brown to black; with thick greyish pruinosity on thorax that is more distinct on scutum; gena, parafacial and lunule brown; inner margin of orbital plate beige; halter whitish-yellow. Calypter yellow, hairs light brown. Genitalia: (Figs. 154–158) Surstylus small, rounded, fused to venter of epandrium, curved posteriorly with dense apical cluster of short setae. Cercus small. Hypandrium broadly rounded; inner lobe V-shaped with two medial setae. Postgonite simple and narrow with one seta. Phallophorus cylindrical with anterodorsal margin lengthened to reach base of basiphallus; lateral lobe simple, narrow. Basiphallus consisting of one pair of long, narrow plates with irregular margins and narrower apices; right plate with base positioned dorsally, left plate with base wrapping around venter. Hypophallus (one pair of lateral sclerites) small, irregular-ovate in outline. Mesophallus not evident. Paraphallus very small, with irregular subovate outline. Distiphallus sharply angled dorsally, thick with apex narrower and slightly curved; seen “ventrally”, outline X-shaped with short arms and long middle section, with apical arms shorter, irregular and less well-defined, each ending in its own small membranous lobe. Ejaculatory apodeme not found. Comments. This is the second agromyzid to be reared from Triosteum, after Aulagromyza orbitalis. Also in Iowa, we have found mines representing two other Phytomyza species, presumably likewise in the periclymeni superspecies. One is an entirely linear mine in the leaf blade, with frass in closely spaced grains and with a brown puparium formed at the end; the other is digitate and centered on the midrib, similar to those of P. chamaemetabola (Griffiths) and P. gregaria Frick on Lonicera involucrata (Richardson) Banks ex Spreng. (Griffiths 1974). Phytomyza triostevena is a modestly sized and almost entirely dark species with one ors and pale greyish-blue pruinosity on the thorax. Among the Caprifoliaceae mining Phytomyza, it can be differentiated by having numerous rows of acrostichal setulae, and a small pair of lateral sclerites on the hypophallus that are irregularly ovate in outline and positioned distal to the end of the basiphallus; this combination of characters will key it approximately to P. nigrilineata (Griffiths) in Griffiths (1974). The small, upturned distiphallus must be examined for confident diagnosis; in lateral view, it is relatively stout with the apex narrowed and curved, and in ventral view, is more than twice as long as wide with shallow bifurcations at either end, appearing somewhat X-shaped; the distal points are shallower, somewhat ill-defined and annulated., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 39-40, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Mews (2019 a) [Photographed leaf mines on orange-fruited horse gentian].","Griffiths, G. C. D. (1974) Studies on boreal Agromyzidae (Diptera). V. On the genus Chromatomyia Hardy, with revision of Caprifoliaceae-mining species. Quaestiones entomologicae, 10, 35 - 69."]}

Phytomyza thermarum (Griffiths) (Fig. 85) Material examined. VERMONT: Windham Co., Marlboro, Hogback Mountain, 42.852158, -72.797828, 21.vi.2018, em. 4.vii.2018, C.S. Eiseman, ex Erigeron pulchellus, # CSE4720, CNC1135661 (1♁). Hosts. Asteraceae: Erigeron philadelphicus L. (Griffiths 1976), E. * pulchellus Michx. Larval mine. (Fig. 85) Partly in the stem, petiole, and basal part of the leaf midrib, with whitish linear channels radiating into the blade. These channels are mostly more or less full-depth but may be on the upper or lower surface. Frass is mostly in discrete grains (partly forming beaded strips in most mines according to Griffiths (1976)). Puparium. White or yellowish-white, formed within the mine, either in the petiole or on the lower surface of the blade, with its anterior spiracles projecting ventrally through the epidermis (Griffiths 1976). Phenology and voltinism. This species is evidently multivoltine in western Canada, with larvae collected in early June emerging as adults within two weeks, larvae and puparia collected in mid-July emerging as adults in late July and early August, and those collected in early September emerging as adults within a month (Griffiths 1976). Distribution. * USA: VT; Canada: AB, BC (Griffiths 1976). Mines possibly representing this species have been found on Erigeron philadelphicus in PA (Schultz 2020)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 38, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Griffiths, G. C. D. (1976) Studies on boreal Agromyzidae (Diptera). XII. Phytomyza and Chromatomyia miners on Astereae (Compositae). Quaestiones entomologicae, 12, 239 - 278.","Schultz, M. (2020) [Photographed leaf mines on Erigeron philadelphicus.] Available from: https: // www. inaturalist. org / observations / 48127531 (accessed 17 July 2020)"]}

Liriomyza euphorbivora Eiseman & Lonsdale, spec. nov. (Figs. 40–43, 75, 140–143) Holotype. USA. OKLAHOMA: Payne Co., Marena, 36.072618°, -97.247242°, 29.v.2018, em. 14.vi.2018, M.W. Palmer, ex Euphorbia marginata, # CSE4617, CNC1144138 (1♁). Etymology. The specific epithet refers to the host plant genus, Euphorbia L. Host. Euphorbiaceae: Euphorbia marginata Pursh. Leaf mine. (Fig. 75) The single collected mine was a whitish (partially greenish), roughly circular blotch on the upper leaf surface, centered on the midrib, 7 mm across when complete, with frass in a few small, dark grains and lumps. A photo taken of the lower leaf surface at the time of collection shows what appears to be a 0.2-mm wide, epidermal linear mine, but this does not appear to be contiguous with the mine that contained the fly larva and we believe it is unrelated. It is not discernible in the preserved leaf. Puparium. Yellow; formed outside the mine. Phenology and voltinism. The only known specimen was collected as a larva in late May, emerging as an adult in mid-June. Distribution. USA: OK. Adult description. Wing length 1.5 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 2.8. Eye height divided by gena height: 3.0. First flagellomere small, rounded. Thorax subshining. Chaetotaxy: Two ori, two ors; subequal to ocellar and postvertical setae. Four dorsocentral setae, only first seta long, with second seta ⅖ length. Five rows of acrostichal setulae. Coloration: (Figs. 40–43) Setae dark brownish-black. Head yellow with ocellar spot dark brown; posterolateral corner of frons dark brown nearly to base of outer vertical, paler brown to base of inner vertical; back of head and clypeus brown. Scutum dark brown with wide, complete lateral yellow stripe on scutum (reaching base of posterior supra-alar and intra-alar) that continues along posterior margin as narrow line along scutellum. Mediotergite dark brown; anatergite brown with posterodorsal corner yellow; katatergite yellow with posteroventral corner brown. Pleuron yellow in base color; anepisternum with wide clavate anteroventral stripe (wider anteriorly) and narrow posteromedial line along suture; anepimeron mottled brown; katepisternum brown below level of seta; meron mostly brown. Scutellum dark anterolaterally. Legs yellow with basal half of fore coxa dark brown, mid coxa basally brownish and hind coxa mottled brown; base of femora narrowly brown (reduced ventrally); tibiae brown with base of fore and mid tibiae yellow, fore tibia yellower and hind tibia darker; tarsi brown. Abdomen brown dorsally, stripe narrowing posteriorly to tergite 5; epandrium brown. Calypter margin yellow with grey tint, hairs greyish-brown. Genitalia: (Figs. 140–143) Epandrium not fused to surstylus, with one spine. Surstylus with two apical spines. Phallophorus with venter much reduced, short. Basiphallus sclerotized along dorsal and left lateral surface with extension on left distal margin below paraphallus. Paraphallus simple, narrow. Hypophallus with small rod-shaped sclerite with apical hairs. Ejaculatory duct swollen and pigmented apically. Mesophallus slightly shorter than distiphallus, subcylindrical, fused to distiphallus dorsally, with slight ventral carina along suture. Distiphallus slightly wider than mesophallus, distal half shallow and cup-shaped, enclosing one pair of short processes, angled dorsally; basal half slightly narrower, sides parallel in ventral view, tapering basally in lateral view. Ejaculatory apodeme large and well-developed with broad blade; sperm pump with broad sclerotized cup-like venter. Comments. This is the second Liriomyza to be reared from Euphorbia in North America, after L. euphorbiella Eiseman & Lonsdale. That species is likewise known only from Payne Co., Oklahoma, but it was reared from E. heterophylla var. cyathophora (Murray) Griseb. rather than E. marginata, and its mine begins with a distinct linear portion and does not cross the midrib. Mines on E. marginata similar to that of L. euphorbivora have been photographed in Ontario (Mews 2019b), and the aborted mine on E. cf. brachycera Engelm. in Colorado noted by Eiseman & Lonsdale (2019) could conceivably have represented this species. TSF has found largely linear Liriomyza mines on E. curtisii Engelm. in North Carolina, but so far adults have not been reared from these. Worldwide, 12 Liriomyza species have now been reported from Euphorbia, including three listed by Benavent- Corai et al. (2005) that were omitted without explanation from the review of Euphorbia -feeding Agromyzidae by Martinez & Sobhian (2000). Of these three, L. congesta (Becker) normally feeds on Fabaceae, and L. sonchi Hendel on Asteraceae, so these records seem likely to be erroneous (both are derived from a paper on parasitoids of Agromyzidae in Valencia, published in 1987). The record of the highly polyphagous L. sativae Blanchard appears plausible, however, apart from its being poorly documented (the cited source, Martinez (1993), is apparently a simple list of host genera, and is a publication we have been unable to obtain after multiple attempts; Martinez & Sobhian (2000) explicitly stated that L. sativae is “not yet recorded on Euphorbia ”). Possibly this record refers to McClanahan (1975), who caged a population of L. sativae with E. pulcherrima Willd. ex Klotzsch and observed light mining but no puparium formation. Whether the unidentified Nearctic mines noted above represent any of these 12 species remains to be seen, but to aid in the identification of Liriomyza specimens reared from Euphorbia, a modified version of the key by Martinez & Sobhian (2000) is presented below with L. euphorbiella, L. euphorbivora, and L. sativae added, and supplemented with data from Ellis (2020). Updated key to the species of Liriomyza feeding on Euphorbia 1. Mesonotum with yellow longitudinal bands................................................................ 2 1′. Mesonotum largely black, without distinct yellow longitudinal bands............................................ 3 2. Back of head, at level of vertical bristles, entirely yellow. Acrostichal setulae less numerous (5 to 8), irregularly arranged in two rows...................................................................... Liriomyza euphorbiae Martinez [mine initially narrow and serpentine, developing into an irregular blotch; on E. kotschyana Fenzl and E. macroclada Boiss.; Turkey] 2′. Back of head, at level of vertical bristles, partly brown. Acrostichal setulae more numerous, arranged in four almost regular rows...................................................................... Liriomyza heringi Nowakowski [mine a blotch; on E. esula L., E. nicaeensis All., E. palustris L., E. salicifolia Host, and possibly E. amygdaloides L.; Lithuania, Romania, former Soviet Union] 3. Wings without vein dm-cu.............................................................................. 4 3′. Wings with vein dm-cu................................................................................. 5 4. Orbits between upper ors and vti completely yellow. Abdominal tergites without yellow lateral band. Penultimate section of M4 2.5 to 3 times longer than last section.............................................. Liriomyza balcanica (Strobl) [mine broadly linear or a linear-blotch; on E. cyparissias L., E. esula, E. myrsinites L., and possibly E. amygdaloides; Germany and Poland to Turkey] 4′. Upper part of orbits black. At least the basal part of abdominal tergite with yellow border. Penultimate section of M4 maximum 2 times longer than last section................................................... Liriomyza myrsinitae Hering [mine a blotch preceded by a short linear portion; on E. myrsinites; Bulgaria, Spain] 5. Orbits between upper ors and vti completely yellow. Mesophallus long, narrow, fused to distiphallus with broad, bilobed cuplike base from which one pair of short weakly scleritized tubules emerge.................. Liriomyza pascuum (Meigen) [mine a primary blotch, generally containing several larvae; on E. amygdaloides, E. characias L., E. dulcis L., E. esula, E. glareosa Pall. ex M.Bieb., E. palustris, and E. pithyusa L.; throughout Europe] 5′. Upper part of orbits black. Phallus not as above............................................................. 6 6. Frons yellow at base of inner vertical seta. Distiphallus entirely split, short, with minute basal stem contrasting rounded apical chamber...................................................................... Liriomyza strigata (Meigen) [branching mine centered on the midrib; a polyphagous species, uncommonly reported from unspecified Euphorbia; throughout Europe] 6′. Frons brown at base of inner vertical seta. If distiphallus entirely split and short (L. huidobrensis), then apical chamber narrow, not much wider than stem............................................................................... 7 7. Surstylus with two apical spines. Distiphallus angled dorsally, with very short apical chamber........................ 8 7′. Surstylus with one apical spine. Distiphallus various, but not as above........................................... 9 8. Eye 4.8–5.9 times higher than gena. Posterior margin of scutum along scutellum black. Ventral 2/3 of anepisternum brown. Femora entirely yellow. Left distal margin of basiphallus long, extending to apex of paraphallus. Basal stem-like half of distiphallus (fused to mesophallus) gradually narrowing to point of fusion with mesophallus............................................................................................ Liriomyza euphorbiella Eiseman & Lonsdale [mine initially narrow and linear with the frass in alternating strips, expanding to an elongate blotch with irregular and more or less diffuse frass; on E. heterophylla var. cyathophora; USA] 8′. Eye 3.0 times higher than gena. Posterior margin of scutum narrowly yellow. Anepisternum yellow with brown clavate spot. Femora brown basally (paler on venter). Left distal margin of basiphallus shorter, not reaching apex of paraphallus. Basal stemlike half of distiphallus parallel-sided and abruptly wider than mesophallus.. Liriomyza euphorbivora Eiseman & Lonsdale [mine a roughly circular blotch on upper leaf surface, centered on the midrib, with frass in a few small, dark grains and lumps; on E. marginata; USA] 9. Femora usually only brown basally, but sometimes more heavily mottled. Abdominal tergites narrowly yellow laterally. Acrostichal setulae (acr) between second and fourth pairs of dorsocentral setae in four more or less regular rows. Distiphallus simple, cup-shaped................................................................... Liriomyza sativae Blanchard [mine linear; a polyphagous species, with one questionable record from unspecified Euphorbia (light mining of E. pulcherrima observed in a laboratory setting, but no puparia were formed); North and South America; introduced in Europe, Asia, Africa, Australia, and several Pacific islands] 9′. Femora brown basally and with additional light to heavy dorsal mottling. Lateral margin of abdominal tergites brown. Acros- tichal setulae (acr) between second and fourth pairs of dorsocentral setae in two to three more or less regular rows. Distiphallus divided into one pair of short tubules......................................... Liriomyza huidobrensis (Blanchard) [mine linear, often following the midrib and lateral veins; a polyphagous species, uncommonly reported from unspecified Euphorbia according to Martinez & Sobhian (2000); the only record listed by Weintraub et al. (2017) is one of E. marginata from China; this South American fly is introduced in North America, Europe, Asia, and Africa] Characters of the thorax of Liriomyza euphorbivora best differentiate it from other Liriomyza externally, as four rows of acrostichal setulae and a very narrow yellow posterior margin on the scutellum is not found in many species. In the United States, it occurs in L. eupatoriella Spencer, but this species has both vertical setae on yellow, the femora are entirely yellow (not basally brown) and the distiphallus is of a different structure, being more cup-shaped with the base strongly narrowed in lateral view. In Canada, the new species keys most readily to L. cracentis Lonsdale and L. agrios Lonsdale, although the spot on the anepisternum of these two species is much larger, their femora are entirely yellow, and the distiphallus is more cup-shaped, without a short apical chamber. The phallus instead shows more similarity to species without the posteriorly yellow scutum. These include L. conclavis Lonsdale (host unknown), L. asclepiadis Spencer and L. peleensis Spencer (both on Asclepias), which have a medially constricted mesophallus and a very sharply upturned distiphallus, and L. subasclepiadis Spencer (also on Asclepias), which also has a more strongly angled distiphallus. Updated key to the species of Liriomyza feeding on Euphorbia 1. Mesonotum with yellow longitudinal bands................................................................ 2 1′. Mesonotum largely black, without distinct yellow longitudinal bands............................................ 3 2. Back of head, at level of vertical bristles, entirely yellow. Acrostichal setulae less numerous (5 to 8), irregularly arranged in two rows...................................................................... Liriomyza euphorbiae Martinez [mine initially narrow and serpentine, developing into an irregular blotch; on E. kotschyana Fenzl and E. macroclada Boiss.; Turkey] 2′. Back of head, at level of vertical bristles, partly brown. Acrostichal setulae more numerous, arranged in four almost regular rows...................................................................... Liriomyza heringi Nowakowski [mine a blotch; on E. esula L., E. nicaeensis All., E. palustris L., E. salicifolia Host, and possibly E. amygdaloides L.; Lithuania, Romania, former Soviet Union] 3. Wings without vein dm-cu.............................................................................. 4 3′. Wings with vein dm-cu................................................................................. 5 4. Orbits between upper ors and vti completely yellow. Abdominal tergites without yellow lateral band. Penultimate section of M4 2.5 to 3 times longer than last section.............................................. Liriomyza balcanica (Strobl) [mine broadly linear or a linear-blotch; on E. cyparissias L., E. esula, E. myrsinites L., and possibly E. amygdaloides; Germany and Poland to Turkey] 4′. Upper part of orbits black. At least the basal part of abdominal tergite with yellow border. Penultimate section of M4 maximum 2 times longer than last section................................................... Liriomyza myrsinitae Hering [mine a blotch preceded by a short linear portion; on E. myrsinites; Bulgaria, Spain] 5. Orbits between upper ors and vti completely yellow. Mesophallus long, narrow, fused to distiphallus with broad, bilobed cuplike base from which one pair of short weakly scleritized tubules emerge.................. Liriomyza pascuum (Meigen) [mine a primary blotch, generally containing several larvae; on E. amygdaloides, E. characias L., E. dulcis L., E. esula, E. glareosa Pall. ex M.Bieb., E. palustris, and E. pithyusa L.; throughout Europe] 5′. Upper part of orbits black. Phallus not as above............................................................. 6 6. Frons yellow at base of inner vertical seta. Distiphallus entirely split, short, with minute basal stem contrasting rounded apical chamber...................................................................... Liriomyza strigata (Meigen) [branching mine centered on the midrib; a polyphagous species, uncommonly reported from unspecified Euphorbia; throughout Europe] 6′. Frons brown at base of inner vertical seta. If distiphallus entirely split and short (L. huidobrensis), then apical chamber narrow, not much wider than stem............................................................................... 7 7. Surstylus with two apical spines. Distiphallus angled dorsally, with very short apical chamber........................ 8 7′. Surstylus with one apical spine. Distiphallus various, but not as above........................................... 9 8. Eye 4.8–5.9 times higher than gena. Posterior margin of scutum along scutellum black. Ventral 2/3 of anepisternum brown. Femora entirely yellow. Left distal margin of basiphallus long, extending to apex of paraphallus. Basal stem-like half of distiphallus (fused to mesophallus) gradually narrowing to point of fusion with mesophallus............................................................................................ Liriomyza euphorbiella Eiseman & Lonsdale [mine initially narrow and linear with the frass in alternating strips, expanding to an elongate blotch with irregular and more or less diffuse frass; on E. heterophylla var. cyathophora; USA] 8′. Eye 3.0 times higher than gena. Posterior margin of scutum narrowly yellow. Anepisternum yellow with brown clavate spot. Femora brown basally (paler on venter). Left distal margin of basiphallus shorter, not reaching apex of paraphallus. Basal stemlike half of distiphallus parallel-sided and abruptly wider than mesophallus.. Liriomyza euphorbivora Eiseman & Lonsdale [mine a roughly circular blotch on upper leaf surface, centered on the midrib, with frass in a few small, dark grains and lumps; on E. marginata; USA] 9. Femora usually only brown basally, but sometimes more heavily mottled. Abdominal tergites narrowly yellow laterally. Acrostichal setulae (acr) between second and fourth pairs of dorsocentral setae in four more or less regular rows. Distiphallus simple, cup-shaped................................................................... Liriomyza sativae Blanchard [mine linear; a polyphagous species, with one questionable record from unspecified Euphorbia (light mining of E. pulcherrima observed in a laboratory setting, but no puparia were formed); North and South America; introduced in Europe, Asia, Africa, Australia, and severa, Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 27-29, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Mews (2019 b) [Photographed leaf mines on snow-on-the-mountain]. Available from: https: // www. inaturalist. org / observations / 34273256 (accessed 15 January 2020)","Martinez, M. & Sobhian, R. (2000) Review of the agromyzid flies associated with Euphorbia spp., and prospects of their use for biological control of leafy spurge in North America. Redia, 81, 1 - 15. [1998]","McClanahan, R. J. (1975) Notes on the vegetable leafminer, Liriomyza sativae (Diptera: Agromyzidae), in Ontario. Proceedings of the Entomological Society of Ontario, 105, 40 - 44. [1974]","Ellis, W. N. (2020) Leafminers and plant galls of Europe. Available from: https: // bladmineerders. nl / (accessed 12 September 2020)","Weintraub, P. G., Scheffer, S. J., Visser, D., Valladares, G., Correa, A. S., Shepard, B. M., Rauf, A., Murphy, S. T., Mujica, N., MacVean, C., Kroschel, J., Kishinevsky, M., Joshi, R. C., Johansen, N. S., Hallett, R. H., Civelek, H. S., Chen, B. & Metzler, H. B. (2017) The invasive Liriomyza huidobrensis (Diptera: Agromyzidae): understanding its pest status and management globally. Journal of Insect Science, 17 (1), 1 - 27. https: // doi. org / 10.1093 / jisesa / iew 121"]}

Phytomyza palmeri Eiseman & Lonsdale Material examined. OKLAHOMA: Payne Co., Mehan, 36.013839°, -96.998103°, 15.xi.2017, em. 22.xi.2017, M.W. Palmer, ex Symphoricarpos orbiculatus, # CSE4380, CNC1135649 (1♁); 30.xii.2017, em. 30.xii.2017, M.W. Palmer, ex Symphoricarpos orbiculatus, # CSE4375, CNC1144114–1144115 (1♁ 1♀). Host. Caprifoliaceae: Symphoricarpos orbiculatus Moench. Leaf mine. Upper surface, gradually widening, at least sometimes linear throughout, but often contorted to form a secondary blotch. The black frass is in closely spaced grains or beaded strips, allowing the larva’s path to be traced even when the mine forms a nearly continuous blotch (Eiseman & Lonsdale 2018). Puparium. Brown; formed within the mine, its anterior spiracles projecting ventrally through the upper epidermis (Eiseman & Lonsdale 2018). A broad, dark ventral stripe is at least sometimes present, as in Phytomyza flavilonicera, but this is not evident in all photographed mines, perhaps due to the rest of the puparium being fairly dark and not providing sufficient contrast to show through the epidermis. Phenology and voltinism. 25 October is the earliest date we have found probable mines of this species, with some mines recently initiated and some already complete. Specimens have been reared from mines collected between 7 November and 30 December, with adults emerging indoors without a pupal diapause (17 November to 30 December). Distribution. USA: OK. Comments. These additional rearings do not provide new host or distribution records for Phytomyza palmeri, but it is worth noting that the genitalia of these specimens differ slightly from those of the type series, representing new intraspecific variation. While agreeing in overall external and genitalic morphology, the apex of the ill-defined and slightly annulated distiphallus of the new specimens is slightly larger, in some cases up to ¼ longer, and one male has a slight medial constriction in the segment (seen ventrally)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 37, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Phytomyza notopleuralis Spencer (Figs. 58, 84) Material examined. MASSACHUSETTS: Berkshire Co., Great Barrington, 42.197870, -73.335897, 17.ix.2017, em. ~ 16.iv.2018, C.S. Eiseman, ex Cornus sericea, # CSE4416, CNC1144083–1144084 (1♁ 1♀). Host. * Cornaceae: Cornus sericea L. Leaf mine. (Fig. 84) Entirely linear, on the upper leaf surface; frass in somewhat liquidy strips along the sides. Puparium. Shining dark brown with paler brown intersegmental boundaries; formed outside the mine. Phenology and voltinism. Our specimens were collected as larvae in mid-September and emerged as adults the following spring. There is presumably at least one earlier generation; the only other known specimens were collected as adults on 3 June (Ontario) and 9 July (Michigan) (Spencer 1969; Spencer & Steyskal 1986). Distribution. USA: *MA, MI; Canada: ON (Spencer & Steyskal 1986). Comments. Neither Spencer (1969) nor Spencer & Steyskal (1986) made any statements about the possible relationships of Phytomyza notopleuralis to other Phytomyza. Winkler et al. (2009) indicated that it is near P. spinaciae Hendel, a Palearctic species mining leaves of Cynareae (Asteraceae) and forming a white puparium within the leaf, and placed it in the syngenesiae group. However, the leaf mine is identical with that of P. agromyzina Meigen, which feeds on various dogwoods including Cornus sericea, and the younger P. notopleuralis may eventually prove to be its junior synonym. External morphology is nearly identical, excluding the presence of a strongly pale yellow notopleuron in P. notopleuralis (Fig. 58) (yellowish tint with margins yellower in P. agromyzina), and the genitalia (Spencer 1969: Fig. 470) only differ in that the distiphallus of P. agromyzina is basally straight (not shallowly curved) and sometimes slightly abbreviated; the apex of the distiphallus is also sometimes a bit straighter. The phallus of P. agromyzina illustrated in Spencer (1969: Fig. 395) shows the distiphallus as a simple membranous lobe without the narrow, apically forked and medially deviated sclerotized band; this is apparently an artifact of illustration, as this structure is present in all dissected material examined at the CNC., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 36-37, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. (1969) The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada, 64, 1 - 311. https: // doi. org / 10.4039 / entm 10164 fv","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","Winkler, I. S., Scheffer, S. J. & Mitter, C. (2009) Molecular phylogeny and systematics of leaf-mining flies (Diptera: Agromyzidae): delimitation of Phytomyza Fallen sensu lato and included species groups, with new insights on morphological and host-use evolution. Systematic Entomology, 34, 260 - 292. [including online Supporting Information] https: // doi. org / 10.1111 / j. 1365 - 3113.2008.00462. x"]}

Cerodontha (Dizygomyza) morosa (Meigen) Material examined. OKLAHOMA: Payne Co., Mehan, 36.013839°, -96.998103°, 20.xii.2017, em. 12.ii.2018, M.W. Palmer, ex Carex amphibola, # CSE4377, CNC1135648 (1♀) Hosts. Cyperaceae: [Carex * amphibola Steud., C. festucacea Willd.], C. gracillima Schwein., C. hitchcockiana Dewey, C. leptonervia (Fernald) Fernald (Eiseman & Lonsdale 2018). Leaf mine. A long, whitish corridor, changing direction a few times, with frass in a single blackish lump toward the leaf apex. Puparium. Yellowish or reddish to blackish-brown; formed within the mine, glued to the floor toward the base of the leaf. Phenology and voltinism. In Massachusetts, Cerodontha morosa has been reared in late June and early July from leaf mines collected in June. In Oklahoma, tentatively identified females have been reared from overwintering pupae, and from mines collected in early and late May with adults emerging in early May and early June (Eiseman & Lonsdale 2018). Distribution. USA: CA, MA, MO (Spencer 1981), [OK]; records from IL, IN, MD, MI (Priest et al. 2020), and SD require verification; Europe (Eiseman & Lonsdale 2018). The Montana record given by Spencer & Steyskal (1986) and repeated by Eiseman & Lonsdale (2018) was evidently an erroneous substitution for the Missouri record given by Spencer (1981). Comments. This is the first record of an agromyzid from Carex amphibola., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 22, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Spencer, K. A. (1981) A revisionary study of the leaf-mining flies (Agromyzidae) of California. University of California Division of Agricultural Sciences Special Publication, 3273, 1 - 489.","Priest, R. J., Kula, R. R. & Gates, M. W. (2020) Leaf mining insects and their parasitoids in the old-growth forest of the Huron Mountains. The Great Lakes Entomologist, 52 (3 - 4), 117 - 160. [2019]","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Phytomyza doellingeriae Eiseman & Lonsdale Material examined. MASSACHUSETTS: Worcester Co., Rutland, 42.381206, -71.964980, 9.vi.2018, em. ~ 29. vi.2018, C.S. Eiseman, ex Doellingeria umbellata, # CSE4705, CNC1135713 (1♁). Host. Asteraceae: Doellingeria umbellata (Mill.) Nees. Leaf mine. Whitish, upper surface, narrow, entirely linear; frass in closely spaced grains or beaded strips along the sides (Eiseman & Lonsdale 2018). Puparium. Blackish; formed outside the mine. Phenology and voltinism. This species is at least bivoltine, with larvae feeding in early June in Massachusetts emerging as adults within a few weeks; in Maine, larvae have been found in late August, with adults emerging the following spring. Distribution. USA: *MA, ME. Comments. Phytomyza solidaginivora Spencer has been reared from larvae forming similar mines on Doellingeria umbellata in mid-June in Maine (Eiseman & Lonsdale 2018); it therefore appears that phenology is not useful in distinguishing mines of the two species., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 34, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Melanagromyza minimoides Spencer (Figs. 62–63) Material examined. IOWA: Winneshiek Co., Decorah, Will Baker Park, 10–13.viii.2017, em. by 16.viii.2017, J. van der Linden, ex Rudbeckia laciniata, # CSE4945, CNC1643663–1643667 (2♁ 3♀). Hosts. Asteraceae: Borrichia frutescens (L.) DC., Helenium flexuosum Raf., Helianthus annuus L., Heliopsis helianthoides Sweet, Melanthera nivea (L.) Small, Rudbeckia laciniata L., Symphyotrichum simmondsii (Small) G.L.Nesom, Verbesina encelioides (Cav.) Benth. & Hook. f. ex A.Gray, V. virginica L.; Cucurbitaceae: Cucurbita foetidissima Kunth; Urticaceae: Urtica dioica L., U. gracilis subsp. holosericea (Nutt.) W.A.Weber (Spencer & Stegmaier 1973; Steyskal 1980b; Spencer & Steyskal 1986; Shi & Gaimari 2015). Larval biology. (Figs. 62–63) On asteraceous hosts, larvae have been consistently reported as feeding in the seedhead. We noted individual disc florets of Rudbeckia laciniata turning brown before the other disc or ray florets had begun to wilt, and opening the affected florets revealed feeding damage to the developing achenes as well as excavations in the receptable. Puparium. (Fig. 63) Whitish-yellow, with black, strongly chitinized posterior spiracular processes, appreciably projecting beyond anal segment (Spencer & Stegmaier 1973); formed in the seedhead. Phenology and voltinism. This species is evidently multivoltine. The Iowa specimens emerged in mid-August, a few days after the puparia were collected. Adults have been reared in Ohio in mid-September (Spencer & Stegmaier 1973), and in Maryland Steyskal (1980b) reported an adult emerging in December (indoors) from host material collected in November. Emergence dates of Florida specimens (reared under unknown conditions) include every month from November to March (Spencer & Stegmaier 1973), and in southern California and Texas adults have been collected in May, with a reared specimen emerging on 1 July (Shi & Gaimari 2015). Distribution. USA: AR, CA, FL, *IA, MD, OH, TX; Mexico; Caribbean; South America (Shi & Gaimari 2015). Comments. Details are lacking for all of the purported rearing records of this species from non-asteraceous hosts. Steyskal (1980b) indicated that a single male (as Melanagromyza radicicola Steyskal) had “emerged indoors from [a] root presumed to be of nettle, Urtica dioica L. ” collected in Maryland. According to Spencer (1990), this specimen was found dead in a rearing container and “[n]o puparium from which it might have emerged could be located. Earlier, specimens of M. minimoides had been reared in the same container from Heliopsis and Rudbeckia.” He thus concluded that the Urtica record was erroneous. Shi & Gaimari (2015) stated that M. minimoides has been reared from U. dioica subsp. holosericea in California, although this is not reflected in their listed label data. Their record from Cucurbita foetidissima is based on a single male labeled “ex. Cucurbita foetidissima (host plant), 1.VII.1963 ”; there is no indication of the larval feeding site or whether a puparium was found., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 10-11, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Stegmaier, C. E. Jr. (1973) Agromyzidae of Florida with a supplement on species from the Caribbean. Arthropods of Florida and Neighboring Land Areas, 7, 1 - 205. https: // doi. org / 10.1007 / 978 - 94 - 017 - 0683 - 4 _ 1","Steyskal, G. C. (1980 b) Six new North American species of Melanagromyza (Diptera, Agromyzidae). Journal of the Washington Academy of Science, 70 (1), 36 - 43.","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","Shi, L. & Gaimari, S. D. (2015) Revision of the genus Melanagromyza in California, with descriptions of three new species (Diptera: Agromyzidae). Zootaxa, 4005 (1), 1 - 102. https: // doi. org / 10.11646 / zootaxa. 4005.1.1","Spencer, K. A. (1990) Host specialization in the world Agromyzidae (Diptera). Series entomologica, 45, i-xii + 1 - 444. https: // doi. org / 10.1007 / 978 - 94 - 009 - 1874 - 0"]}

Liriomyza hypopolymnia Eiseman & Lonsdale, spec. nov. (Figs. 44–47, 76–77, 144–147) Holotype. USA: IOWA: Winneshiek Co., Craggy Rocks, 43°25’58.82”N, 92° 0’29.92”W, 20.x.2017, em. 18.iv.2018, MJ Hatfield, ex Polymnia canadensis, CNC 1144061 (1♁). Paratypes. USA: IOWA: same data as holotype, CNC1144055, CNC1144058, CNC1144062 (2♁ 1♀), Deco- rah, Dug Road, 30.vi.2017, em. by 21.vii.2017, J. van der Linden, ex Polymnia canadensis, # CSE4654, CNC1144029 (1♀). Additional material examined. same data as holotype, CNC1144054, CNC1144056–1144057, CNC1144065– 1144073 (13 puparia). Etymology. The specific epithet refers to the position of the larval mines (Gr. hypo —“under, beneath”) on the lower surface of leaves of Polymnia Kalm. Host. Asteraceae: Polymnia canadensis L. Leaf mine. (Figs. 76–77) A long, rather indistinct, linear mine on the lower leaf surface, with frass in grains and beaded strips along the sides; the mine may be evident from above as a faint green line or mottling. Some mines intermittently switch to the upper surface, where they are much more distinct and appear whitish. Puparium. (Fig. 47) Yellow; formed outside the mine. Phenology and voltinism. This species is at least bivoltine in Iowa. Larvae have been found in late June, with an adult emerging within a few weeks; larvae of the overwintering generation have been found from mid-October to mid-November, emerging as adults in spring. Distribution. USA: IA; leaf mines have also been found in PA (Woods 2018) and TN. Adult description. Wing length 1.9–2.0 mm (♁), 1.9 mm (♀). Length of ultimate section of vein M 4 divided by penultimate section: 2.5–3.1. Eye height divided by gena height: 5.3–7.0. First flagellomere rounded and slightly enlarged, being higher than long; distal margin with longer hairs subequal to width of base of arista. Thorax subshining. Vein dm-m slightly angled. Chaetotaxy: Two ors, two ori; anterior ori slightly shorter. Four dorsocentral setae, decreasing in length anteriorly, only posterior two pairs strong. Six rows of acrostichal setulae; reduced in number posteriorly, typical of Liriomyza, but posterior-most pairs between first pair of dorsocentrals atypical in being widely spaced and incurved. Eye bare and mid tibia without medial setae (eye often minutely haired and mid tibia with posteromedial seta in Liriomyza with convergent posteromedial acrostichal setulae). Coloration: (Figs. 44–46) Overall color of most specimens relatively pale yellow compared to congeners, perhaps because specimens were preserved while still slightly teneral; base color of most whitish-yellow with pigmented patches brown to orange-brown, but female specimen CSE4654 with color typical of most other Liriomyza with pigment dark brown. Setae brown with yellowish shine (shine not pronounced on major setae of thorax and head). Ocellar tubercle light brown; posterolateral corner of frons with light brown spot reaching margin of eye, but not base of vertical setae; back of head brown with venter yellow. Scutum brown with complete lateral yellow stripe that is relatively wide and encompasses base of posterior inter-alar and supra-alar setae; posterior margin of scutum yellow with quadrate spot between dorsocentral rows that has anterior corners extending as one pair of small short points approaching bases of second dorsocentrals; spot on postpronotum and anterolateral spot on scutellum faded, small. Mediotergite dark brown; anatergite with small brown anteroventral spot extending onto posteroventral corner of katatergite. Anepisternum with very small faded anteroventral spot; anepimeron entirely pale; katepisternum brown ventrally, far from base of seta; meron brown on ventral half. Hind tarsus sometimes brownish apically. Abdomen paler brown dorsally with wide yellow lateral margin; stripe narrows to medial spot on tergite 5; epandrium light brown with dorsomedial region light yellow. Calypter margin slightly greyish, hairs yellow. Genitalia: (Figs. 144–147) Epandrium separate from surstylus, with one pair of small ventral spines. Surstylus with one pair of large spines (apical and basal). Phallophorus with venter reduced, short. Basiphallus sclerotized along dorsal and left lateral surfaces, with narrow process extending from left apical margin. Hypophallus small, narrow, with short apical hairs. Paraphallus very small, narrow. Ejaculatory duct swollen and pigmented apically. Mesophallus subcylindrical, tapered at base and apex, very lightly fused to distiphallus; venter flared along suture. Distiphallus wider and longer than mesophallus, cup-shaped with base tapered (abruptly constricted in ventral view), widest before apex, enclosing small internal processes. Ejaculatory apodeme very pale, narrow; sperm pump with small ventral sclerotization. Comments. The only previous record of an agromyzid from Polymnia is that of an undetermined stem-boring Melanagromyza species we reared from P. canadensis in Iowa (Eiseman & Lonsdale 2019). The host of the Venezuelan M. polymniae Spencer is Smallanthus riparius (Kunth) H.Rob. (Spencer 1990). Liriomyza hypopolymnia is distinct in being quite pale with the vertical setae on yellow, the abdomen largely yellow and the scutum posteriorly yellow with a subquadrate yellow spot in front of the scutellum between the dorsocentral rows. This subquadrate spot has small points anterolaterally along the dorsocentral rows, and the few acrostichal setulae within the spot are incurved. These characters are also seen in some L. blechi, except this species usually has the brown scutal spot divided into bands, the third dorsocentral is most reduced (not with anterior two pairs of dorsocentrals reduced with the fourth smallest), and the mid tibia has two posteromedial setae. The male genitalia of this species are also drastically different (see Lonsdale 2017a: Figs. 29–32). The genitalia of L. hypopolymnia more closely resemble those of other species with a large, pale cup-shaped distiphallus such as L. eupatorii, but the new species differs from most of these in having a lateromedially constricted distiphallus and a surstylus with two spines (one apical and one basal); both of these features are shared by L. sabaziae Spencer, but this is a much darker species with a slightly smaller distiphallus that is dorsoventrally compressed on the basal half (see Lonsdale 2011: Figs. 202, 203)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 29-30, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Woods, P. (2018) Linear mines in Polymnia canadensis. Available from: https: // bugguide. net / node / view / 1489388 (accessed 13 December 2019)","Spencer, K. A. (1990) Host specialization in the world Agromyzidae (Diptera). Series entomologica, 45, i-xii + 1 - 444. https: // doi. org / 10.1007 / 978 - 94 - 009 - 1874 - 0","Lonsdale, O. (2017 a) The Liriomyza (Agromyzidae: Schizophora: Diptera) of Canada & Alaska. Zootaxa, 4234 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4234.1.1","Lonsdale, O. (2011) The Liriomyza (Agromyzidae: Schizophora: Diptera) of California. Zootaxa, 2850 (1), 1 - 123. https: // doi. org / 10.11646 / zootaxa. 2850.1.1"]}

Agromyza pseudoreptans Nowakowski Material examined. MASSACHUSETTS: Franklin Co., Northfield, 276 Old Wendell Rd., 26.x–1.xi.2017, em. 2–8.v.2018, C.S. Eiseman, ex Urtica dioica, # CSE4463, CNC1135695–1135702 (3♁ 5♀). Photographed mines. IOWA: Linn Co., Cedar Rapids / ICNC, 6.xi.2011, J. Zito, Urtica dioica [vacated mine], BG 592789; MARYLAND: Harford Co., Susquehanna State Park, 28.v.2018, J. Emm, Urtica dioica, BG 1527598. Host. Urticaceae: Urtica dioica L. Leaf mine. A dark greenish linear-blotch, turning brown; following the leaf margin for several serrations and then expanding away from it. The egg is often laid near the leaf margin but may be laid along a lateral vein away from the margin, in which case the initial linear portion meanders until reaching the margin. Frass is deposited in numerous irregular, dark lumps. Puparium. Brown; formed outside the mine. Phenology and voltinism. This species is bivoltine, with a summer pupal diapause and an exceptionally latefeeding fall generation. A photograph taken in Maryland shows an occupied mine on 28 May. A larva collected in Vermont in late June emerged as an adult in mid-October (Eiseman & Lonsdale 2018). Larvae of our new reared series were active in Massachusetts from late October into early November, and in Washington we found larvae feeding as late as 3 December 2018. Distribution. USA: AK, CA, [IA], ID, *MA, [MD], MN, OR, VT, WA; Canada: AB, BC, MB, NT, ON, QC; Europe; Georgia, Kazakhstan; Russian Far East (Nartshuk & von Tschirnhaus 2017; Eiseman & Lonsdale 2018)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 6-7, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Nartshuk, E. & von Tschirnhaus, M. (2017) Leafminer flies (Diptera: Agromyzidae) of the fauna of Russia and adjacent countries: The genus Agromyza Fallen. Studia dipterologica, 22 (2), 215 - 232. [2015]"]}

Melanagromyza hieracii Eiseman & Lonsdale, spec. nov. (Figs. 9–12, 61, 99–104) Holotype. USA. IOWA: Allamakee Co., Fish Farm Mounds WMA, 30.ix.2017, em. spring 2018, J. van der Linden, ex Hieracium scabrum, # CSE4669, CNC1144040 (1♁). Etymology. The specific epithet refers to the host plant genus, Hieracium L. Host. Asteraceae: Hieracium scabrum Michx. Larval biology. (Fig. 61) The larva bores in the pith of the stem, forming a linear gallery. Puparium. (Fig. 12) Straw-colored, formed at the end of the larval gallery; rear spiracular horns pronounced, blackish, curved and pointed. Phenology and voltinism. The holotype emerged after overwintering from a puparium collected on 30 September. All other puparia found at that time in stems of H. scabrum were already empty. Assuming these represented the same species, this suggests either that Melanagromyza hieracii has more than one generation or that adults may be capable of overwintering. Distribution. USA: IA. Adult description. Wing length 2.7 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 0.7. Eye height divided by gena height: 4.5. First flagellomere small, rounded. Ocellar triangle largely obscured by detritus, appearing to reach past posterior ori. Anterior half of gena slightly angled upwards. Cheek not evident. Clypeus rounded. Eye and head longest above midpoint. Thorax subshining. Body in relatively poor condition; partially collapsed, some setae missing. Chaetotaxy: Four gracile ori strongly incurved (nearly flat on frons), two strong ors; setae long. Several rows of scattered orbital setulae, as long as eye hairs, erect (inner setulae) to reclinate (outer). Ocellar setae possibly missing, bases obscured. Eye hairs sparse and scattered on dorsal half with dense dorsomedial patch. Anterior of two dorsocentral setae missing, sockets large. Acrostichal setulae in ten irregular rows. Katepisternum with one additional subdominant seta. Mid tibia with two posteromedial setae. Coloration: (Figs. 9–11) Setae dark brown. Body dark brown, including halter; notum faintly green metallic; abdomen green metallic with blue tint; wing veins brown. Calypter margin and hairs yellow. Genitalia: (Figs. 99–104) Epandrium with posteroventral spine. Surstylus fused to anteroventral margin of epandrium, relatively narrow and produced with slight posterodistal extension; distal margin with irregular rows of tubercle-like setae. Cercus large and well-developed. Hypandrium broadly rounded basally, with long narrow apical apodeme; inner lobe U-shaped with minute setulae. Phallophorus swollen ventrally, base much narrowed. Basiphallus U-shaped (right side irregularly sclerotized in dissected male), with short gap between apex and mesophallus. Mesophallus swollen cylindrical, narrowest at point of insertion to distiphallus, which is relatively posterobasal. Distiphallus short, globular, with short ventromedial swelling into which mesophallus inserts; basal section mostly made up of posteriorly convergent tubules flanking mesophallus; ventral plate subquadrate, distal margin slightly flared laterally; dorsal chamber shifted distally, widest subapically, with one pair of minutely spinulose internal pads; tubular process narrow, distal half exposed, sinuate when viewed laterally. Ejaculatory apodeme with short dark stem with long lateromedial process; blade narrow with medial rib, margin along one side narrow and irregular (broken?), other side wider and rounded with margin clear; sperm pump with dark transverse bar upcurved at ends. Comments. This is the first record of a North American Melanagromyza from Hieracium. The European M. oligophaga Spencer includes H. umbellatum L. among its hosts (Spencer 1990). Externally, Melanagromyza hieracii is not unlike many other congeners with a metallic green shine (weak on the thorax and with a blue tint on the abdomen), a white calypter, two dorsocentrals and a shallow orbital plate. The key in Spencer & Steyskal (1986) will bring the species to M. longensis Spencer / M. virginiensis Spencer, but unlike these species, it has four ori. The phallus is unlike those of these two species in that the dorsal chamber of the distiphallus is parallel-sided, the lateral margins of the distiphallus are very strongly flared apically, the medial tubule is pronounced and long, far exceeding the apex of the short dorsal chamber, and the mesophallus is strongly shifted basally on the distiphallus. The combination of these phallic features, especially the flared distolateral margins of the distiphallus and the basally shifted mesophallus, is quite unusual for the genus. Also see comments for Melanagromyza urticae.

Aulagromyza luteoscutellata (de Meijere) Material examined. MASSACHUSETTS: Nantucket Co., Nantucket, Dead Horse Valley, 6.xi.2017, em. 26– 30.iv.2018, C.S. Eiseman, ex Lonicera morrowii, # CSE4440, CNC1135691–1135692 (1♁ 1♀). Hosts. Caprifoliaceae: [Lonicera canadensis J.Bartram & W.Bartram ex Marshall], L. morrowii A.Gray, L. tatarica L., [Symphoricarpos albus (L.) S.F.Blake] (Eiseman & Lonsdale 2018). Leaf mine. Widening, linear, sometimes becoming contorted; filled centrally with diffuse green frass. In some cases the egg is inserted near the leaf margin and the larva at first follows the margin toward the apex, but mines are frequently formed away from the leaf margin. Puparium. Yellow to yellowish-brown; formed outside the mine. Phenology and voltinism. Previous North American records of this species have documented larvae feeding in May and June in New York and Massachusetts (Eiseman & Lonsdale 2018; Scheffer & Lonsdale 2018), and early July in Ontario (Spencer 1969), with adults emerging within a month in each case. Spencer & Steyskal (1986) gave a date of 13 September for specimens reared in Wisconsin, but it is unclear whether this was the collection or emergence date. Our new specimens from Massachusetts were collected as larvae in early November, overwintered as pupae, and emerged as adults about a month after removal from refrigeration. Distribution. USA: MA, [MN], NY, WI; Canada: AB?, ON, QC; Europe (Eiseman & Lonsdale 2018). Comments. Although the new specimens do not represent a new host or distribution record, the presence of larvae actively feeding in early November is noteworthy. Agromyza pseudoreptans (discussed above) is the only other agromyzid we can recall actively mining leaves in New England as late as November. The only hosts of Aulagromyza luteoscutellata confirmed by rearing in North America are invasive honeysuckles (Lonicera spp.) introduced from the Palearctic. Further rearing from native Caprifoliaceae will be needed to clarify the host ranges and leaf mine characteristics of Aulagromyza known or suspected to feed on these plants, which include seven described species and two undescribed (known only from females)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 20-21, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Scheffer, S. J. & Lonsdale, O. (2018) A survey of Agromyzidae (Diptera) reared from leafmines on Long Island, New York; host associations, distribution data, and the description and host association of a new species. Zootaxa, 4450 (1), 77 - 90. https: // doi. org / 10.11646 / zootaxa. 4450.1.5","Spencer, K. A. (1969) The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada, 64, 1 - 311. https: // doi. org / 10.4039 / entm 10164 fv","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}

Phytomyza confusa Eiseman & Lonsdale Material examined. IOWA: Winneshiek Co., Beard Farm, 23.v.2017, em.?, J. van der Linden, ex Hydrophyllum virginianum, # CSE4645, CNC1135657 (1♁). Host. Hydrophyllaceae: Hydrophyllum virginianum L. Leaf mine. Irregular in shape, sometimes with a distinct linear portion and sometimes seeming to form a primary blotch, even when very small. Mines are whitish with brownish patches and discrete, dark frass grains (Eiseman & Lonsdale 2018). Puparium. Pale brown; apparently formed externally, but this requires confirmation (Eiseman & Lonsdale 2018). Phenology and voltinism. Larvae have now been found in May and September (the latter overwintering as pupae), indicating Phytomyza confusa is at least bivoltine. Distribution. USA: IA. Comments. Although no details of the leaf mine characteristics or pupation site were recorded for this new rearing, it confirms the tentative identification of the host plant that was given in the original description., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 33-34, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Phytomyza cf. aesculi Eiseman & Lonsdale (Fig. 81) Material examined. VIRGINIA: Radford, Wildwood Park, 3.v.2017, em. 8–11.iv.2018, N. V. Kent, ex Aesculus flava, # CSE4400, CNC1135662–1135664 (3♀). Hosts. The type series of Phytomyza aesculi was reared from Sapindaceae: Aesculus glabra Willd. (Eiseman & Lonsdale 2018). The Virginia specimens were reared from A. flava Sol., and mines have also been found on A. pavia L. and A. sylvatica W.Bartram. Leaf mine. (Fig. 81) Upper-surface; mines associated with the type series were described as “linear, gradually widening from 0.3 mm to 2–3 mm; whitish, with liquidy frass forming a broad, green central band, and with black- ish particles scattered within this.” The mines on Aesculus flava are initially narrowly linear with frass in closely spaced grains along the sides, but then rather quickly widen to an elongate blotch ~ 4.5 mm wide, in which the frass is scattered at random. The completed mine is thus shorter and more compact than the mines on A. glabra. On both hosts, there are often multiple mines per leaflet. Puparium. Yellowish-brown to reddish-brown; on A. flava, frequently formed in the exit slit at the end of the mine, or adhering to the upper leaf surface nearby. Phenology and voltinism. Phytomyza aesculi is univoltine, with larvae feeding in spring and adults emerging the following spring. The type series was collected as larvae in central Ohio between 6 and 11 May. A photo on iNaturalist shows larvae in St. Louis, Missouri on 24 April; photos on BugGuide. net show puparia in Radford, Virginia as early as 1 May, and we have found mines as early as 26 March in North Carolina. Distribution. USA: OH; see Comments. Comments. Confirmed specimens of Phytomyza aesculi are only known from USA: OH; leaf mines on Aesculus glabra and undetermined Aesculus spp. matching those of the type series have been found in IA, KS, MN, MO, and Canada: ON. Some mines on A. pavia and A. sylvatica from AL, AR, GA, NC, and SC are long and narrow like those from OH, while others become blotchy like those from which the VA specimens were reared, as do those found on A. glabra in PA. It is possible that there is a single Phytomyza species on Aesculus with somewhat variable leaf mine morphology, but the VA females differ from the type series of P. aesculi in having the face darker, the venter of the lunule and the region around the antennal bases blackish, and the apices of the mid and hind femora are dark. More reared males will be needed to clarify the situation., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 32-33, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Ophiomyia rugula Eiseman & Lonsdale, spec. nov. (Figs. 29–31, 70–72, 127–132) Holotype. USA. NORTH CAROLINA: Durham Co., Durham, Leigh Farm Park, 29.iv.2018, em. 17.v.2018, T. S. Feldman, ex Baccharis halimifolia, # CSE4553, CNC1135680 (1♁). Paratype. Same data as holotype, CNC1135681 (1♁); Wake Co., Morrisville, Lake Crabtree County Park, 12.vii.2018, em. 13.vii.2018, T. S. Feldman, ex Baccharis halimifolia, # CSE4951, CNC934530 (1♁). Etymology. The specific epithet (L. rugula —a small wrinkle) refers to the appearance of the larval feeding site as a small wrinkle in the leaf. Host. Asteraceae: Baccharis halimifolia L. Leaf mine. (Figs. 70–72) Mostly green and interparenchymal, consisting of a short, narrow wrinkle along the midrib. In some examples the mine is confined to the basal portion of the midrib, and in others the basal portion appears to have been mined but the mine is only conspicuously wide or wrinkled at or somewhat beyond the middle of the leaf. Puparium. (Fig. 72) White; centered on the midrib in the distal end of the mine. Phenology and voltinism. This species is evidently at least bivoltine, with larvae feeding in April emerging as adults in mid-May, and with adults of another generation emerging in mid-July. Distribution. USA: NC. Adult description. Wing length 2.3–2.4 mm (♁). Female unknown. Length of ultimate section of vein M 4 divided by penultimate section: 0.8. Eye height divided by gena height: 6.2–6.5. First flagellomere subcircular with slightly longer hairs on anterodorsal margin. Orbital plate very narrow. Ocellar triangle slightly shinier, ending near level of anterior ors. Face with shallow, rounded ridge. Gena rounded anteriorly, only slightly extending past anterior margin of eye; neither pointed nor projecting. Lunule somewhat semicircular with wide medial groove. Palpus narrowed basally. Clypeus with narrow, truncated apical margin with sharp corners. Thorax subshining. Chaetotaxy: One ori, three ors; anterior ors slightly shorter than other fronto-orbitals; paired setae not equally positioned on both sides. Single row of orbital setulae. Ocellar and postvertical setae subequal to posterior ors. Two strong dorsocentral setae. Eight rows of acrostichal setulae. Katepisternum with several enlarged dorsal setae, including one nearly as long as dominant seta. Mid tibia with two posteromedial setae. Coloration: (Figs. 29–31) Setae dark brown. Body dark brown with halter stem light brown with dark patches. Calypter margin and hairs dark brown. Genitalia: (Figs. 127–132) Surstylus broad, rounded, fused to epandrium, with short, pointed tubercle-like setae along inner-distal margin. Cercus large, well-developed. Hypandrium stout medially; inner lobe V-shaped, nearly smooth. Phallophorus cylindrical, weakly connected dorsally to basiphallus; phallus past phallophorus angled ventrally. Basiphallus with short, wide, thin dorsal sclerite, laterally produced into one pair of long sinuate ventral bands and one pair of more dorsally positioned pockets. Mesophallus lightly sclerotized, straight, resembling slightly thickened terminal section of ejaculatory duct. Distiphallus band-like, dorsoventrally compressed, slightly wider than mesophallus (both weakly differentiated in ventral view), which is inserted medially on ventral surface. Additional small U-shaped sclerite suspended in membrane ventral to apex of mesophallus of uncertain homology. Comments. Ophiomyia rugula is clearly related to O. similata (Malloch) based on genitalic structure, and is indistinguishable from it externally, although O. similata has one or two mid tibial setae and most of the type series has smaller fronto-orbitals that are less than half the width of the frons (not slightly more). The genitalia of O. similata differ in lacking the ventrolateral bands of the basiphallus, the distiphallus is wider and higher, and the suspended ventrodistal sclerite of uncertain origin is thicker and more distally positioned. The immature stages of Ophiomyia similata are unknown, but O. abutilivora (discussed above) belongs to a related lineage, and it forms a characteristic gall-like raised mine or “welt” in the stem of Abutilon theophrasti (Malvaceae). Ophiomyia tiliae (Couden) also belongs to this lineage, and it forms galls in twigs of Tilia americana L. (Malvaceae) (Spencer & Steyskal 1986). The feeding damage of O. rugula is also gall-like, and it may be that O. similata and allied species of the California O. jacintensis group (see Spencer & Steyskal (1986: 37)) will prove to have similar larval biology. The only Ophiomyia previously associated with Baccharis is O. spicatae Spencer, known from a single female reared from a stem mine on B. spicata (Lam.) Baill. in Brazil (Spencer 1963)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 19-20, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606","Spencer, K. A. (1963) A synopsis of the Neotropical Agromyzidae (Diptera). Transactions of the Royal Entomological Society of London, 115 (12), 291 - 389. https: // doi. org / 10.1111 / j. 1365 - 2311.1963. tb 00811. x"]}

Aulagromyza orbitalis (Melander) Material examined. OKLAHOMA: Payne Co., Mehan, 36.013839°, -96.998103°, 24.iv.2017, em. 22.iv.2018, M.W. Palmer, ex Symphoricarpos orbiculatus, # CSE4424, CNC1144117 (1♀); WASHINGTON: Thurston Co., Olympia, Centralia Western Trail at 45th Ave., 29.ix.2017, em. 19.ii.2018, E. Stansbury, ex Symphoricarpos albus, # CSE4606, CNC1135596 (1♁). Hosts. Caprifoliaceae: Diervilla lonicera Mill., Lonicera dioica L., L. sempervirens L., Symphoricarpos albus (L.) S.F.Blake, S. mollis Nutt., S. * orbiculatus Moench, S. rotundifolius A.Gray., Triosteum aurantiacum E.P.Bicknell; a mine possibly of this species found on T. perfoliatum L. (Eiseman & Lonsdale 2018). Leaf mine. Variously placed on the upper surface; green to yellowish-green or whitish; in some cases linear throughout, in others widening to a more or less distinct blotch (Eiseman & Lonsdale 2018). On some hosts the frass tends to be regularly deposited in alternating, closely spaced particles or streaks in the narrow linear portion, but on Symphoricarpos orbiculatus it is indistinct or diffuse throughout. Puparium. Yellowish-brown to brown; formed outside the mine. Phenology and voltinism. Our Oklahoma females were collected as larvae in late April and emerged as adults the following spring; a full-year diapause was also reported by Spencer (1969) for a female collected as a larva in Alberta in early June. Otherwise, adults have emerged within a few weeks from larvae collected in North Carolina in April, and from Alberta, Iowa, and Massachusetts in June (Eiseman & Lonsdale 2018). Our new Washington specimens were collected as larvae in late September, emerging after overwintering. Distribution. USA: CA, IA, ID, KS, MA, MI, NC, *OK, WA; Canada: AB, MB (Eiseman & Lonsdale 2018)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on page 21, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Spencer, K. A. (1969) The Agromyzidae of Canada and Alaska. Memoirs of the Entomological Society of Canada, 64, 1 - 311. https: // doi. org / 10.4039 / entm 10164 fv"]}

Liriomyza orilliensis Spencer (Fig. 78) Material examined. IOWA: Winneshiek Co., Decorah, Twin Springs Park, 24.viii.2017, em.?, J. van der Linden, ex Prenanthes alba, # CSE4644, CNC1144028 (1♀). Photographed mines. MASSACHUSETTS: Franklin Co., Northfield, 276 Old Wendell Rd., 7.x.2015, C.S. Eiseman, Nabalus altissimus [aborted/predated mines]; Nantucket Co., Squam Swamp, 8.ix.2019, C.S. Eiseman, Nabalus trifoliolatus; MICHIGAN: Ingham Co., Okemos, Ted Black Woods, 19.vi.2019, C.S. Eiseman, Nabalus; NORTH CAROLINA: Guilford Co., Browns Summit, Haw River State Park, 29.v.2015, T.S. Feldman, Prenanthes alba [vacated mine], BG 1075103; VERMONT: Chittenden Co., Williston, Mud Pond, 28.viii.2016, C.S. Eiseman, Nabalus [vacated mines]. Hosts. Asteraceae: Nabalus * albus (L.) Hook., [N. altissimus (L.) Hook., N. trifoliolatus Cass.], Taraxacum F.H.Wigg. (Eiseman & Lonsdale 2018). Leaf mine. (Fig. 78) Pale greenish to whitish, turning brownish; in some cases beginning as an irregular track, gradually expanding to an elongate blotch, with frass mostly in squiggly strips; in others, forming a more or less compact blotch with distinct secondary feeding lines, and with frass mostly in fine grains. Two or three larvae may feed together. Puparium. Yellow to brown; formed outside the mine. Phenology and voltinism. Data from the few known specimens of this species indicate it is at least bivoltine. Larvae collected in Massachusetts in mid-June emerged as adults in early July (Eiseman & Lonsdale 2018); a larva collected in Alberta in early August emerged as an adult the following spring (Lonsdale 2017a); and the Iowa specimen emerged on an unknown date from a larva collected in late August. Distribution. USA: *IA, MA, [MI, NC, VT]; Canada: AB, ON (Eiseman & Lonsdale 2018)., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 30-31, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Lonsdale, O. (2017 a) The Liriomyza (Agromyzidae: Schizophora: Diptera) of Canada & Alaska. Zootaxa, 4234 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4234.1.1"]}

Haplopeodes loprestii Eiseman & Lonsdale, spec. nov. (Figs. 32–39, 133–139) Holotype. USA. CALIFORNIA: San Diego Co., Coyote Canyon, 14.iii.2018, em. 18.v–3.vi.2018, E. LoPresti, ex Fagonia laevis, # CSE4545, CNC1135597 (1♁). Paratype. Same data as holotype, CNC1135598 (1♀ ) . Etymology. This species is named for Eric F. LoPresti, ecologist and evolutionary biologist, who collected the host material from which it was reared. Host. Zygophyllaceae: Fagonia laevis Standl. Larval biology. Apparently a leafminer. The host material was collected to rear leaf-mining larvae of a gelechiid moth, and the presence of the flies only became evident when the puparia appeared. The rearing vial also contained fruits and stems, so the possibility of these as feeding sites cannot be entirely excluded, but as far as is known all Haplopeodes species are leafminers. Puparium. (Fig. 34) Pale, straw-colored; formed externally. Phenology and voltinism. The two known specimens were reared from larvae that fed in March, pupating by the 20 th and emerging as adults four to six weeks later. Distribution. USA: CA. Adult description. Wing length 1.2 mm (♁), 1.4 mm (♀). Vein dm-m absent. Eye height divided by gena height: 3.2–3.5. First flagellomere slightly longer than high, apically rounded. Arista short, only 1.5 times length of first flagellomere. Orbital plate shallow but distinctly projecting when viewed laterally, especially anteriorly on head; continuing as modestly developed ring around eye. Seen dorsally, frons wider than eye, slightly broader anteriorly. Anterior ocellus slightly displaced anteriorly. Epistoma large, broad, subrectangular, resulting in correspondingly shorter face; face with shallow medial carina. Palpus small, narrow, length approximately three times width. Thorax subshining. Wing veins costalized (closely spaced anteriorly), with width of cell r 4+5 subequal to that of cell sc; length of costal cell subequal to length of cell r 1 past insertion of vein R 1. Chaetotaxy: Setae short. Three ori, very gracile; ors absent. Few orbital setulae, slightly inclinate and erect, partially proclinate. Ocellar seta straight, approximately ¾ length of ori. Postvertical seta absent. Five dorsocentral setae, strongly decreasing in length anteriorly; female with additional small setula in dorsocentral row anteriorly. Only one notopleural seta, medially positioned. Notal setulae mostly absent; female with 1 anteromedial acrostichal setula and with few setulae in intra-alar region. One pair of lateral scutellar setae and two pairs of apical scutellar setae (left anterior seta absent in male). Katepisternal seta not visible in male. Coloration: (Figs. 32–33, 35–39) Setae brown to dark brown with pale brown shine. Base color light yellow with head paler. First flagellomere brown with inner surface (not including margins) yellow; ocellar triangle dark brown; vertex light yellow; posterolateral corner of frons with brown patch fading to base of inner vertical seta; face deeper yellow with slight orange tint; venter of gena with brownish line; back of head dark brown with dorsomedial region paler and venter yellow. Notum dark brown to brown; center of scutellum and mediotergite slightly paler brown; postpronotum light brown, becoming yellower anteriorly and laterally; notopleuron and supra-alar spot light brown to yellow, with notopleuron yellower medially; katatergite light brown. Proepisternum light brown; anepisternum with anteroventral corner paler yellow, bordered posteriorly by oblique brown stripe; katepisternum with brown spot below level of seta; anepimeron irregularly brown; meron mostly dark brown. Legs light yellow; base of coxae narrowly brown; femora with indistinct brownish tint on apical ¼–½; tibiae brown with mediolateral and medioventral regions broadly yellow; tarsi brown. Abdomen brown with lateral margin of tergites narrowly yellow; epandrium darker brown. Calypter margin and hairs yellow. Variation. Female: Darker than male. Setae darker brown to blackish. Only inner-basal surface of first flagellomere yellow; scape and pedicel yellow. Posterolateral corner of frons darker. Paler parts of scutum brown tinted. Anepisternum, anepimeron and meron brown; base color of katepisternum brownish-orange. Femora extensively brown mottled. Abdominal tergites entirely brown. Genitalia: (Figs. 133–139) Epandrium shallow. Surstylus short, incurved, apical margin irregular but essentially straight, without tubercle-like setae or narrow protrusions referred to as “teeth” in Steyskal (1980a); base (concealed within epandrium) incurved. Cerci weakly sclerotized, much thicker laterally, closely spaced. Hypandrium with short, semicircular arch; inner lobe weakly sclerotized, more so on distal margin bearing single seta; arm long, well-defined. Postgonite dark, narrow, with single subapical seta. Phallus small and weakly sclerotized, but comparatively large and dark for Haplopeodes; phallophorus cylindrical, fused to curved basiphallus, which is represented by short ventral sclerotization that is slightly elongated on left side. Ejaculatory duct strongly upcurved past basiphallus, with exposed portion approximately as long as basiphallus; apex rounded, dorsoventrally flattened and lightly sclerotized. Ejaculatory apodeme narrow but well-developed, darker stem grading into blade. Comments. This is the first record of any agromyzid from Fagonia, and the only agromyzids previously recorded from Zygophyllaceae are extremely polyphagous Liriomyza spp. Known hosts of Haplopeodes are otherwise restricted to Amaranthaceae, Portulacaceae, and Solanaceae (Benavent-Corai et al. 2005). While minor color differences between the sexes are evident in some Haplopeodes, the sexual dimorphism observed here is pronounced and unique, with the female being much darker than the male. Both sexes are considerably darker than other Haplopeodes, with the scutellum entirely dark brown (unique among Nearctic Haplopeodes), the first flagellomere and scutellum brown, the pale grey pruinosity usually observed on the thorax is absent (the scutum is subshining), the tibiae are brown with yellow medial regions and the femora are brown mottled. Furthermore, the acrostichal setulae are virtually absent, the orbital plate and parafacial are produced, an epistoma is present, the radial veins are crowded anteriorly, the phallus is pigmented apically and surstylus spines are absent.

Phytomyza tiarellae Griffiths Material examined. WASHINGTON: Thurston Co., Lacey, Chehalis Western Trail / S. 45th Ave., 11.iii.2018, em.?, E. Stansbury, ex Tolmiea menziesii, # CSE4608, CNC1135594–1135595 (1♁ 1♀); Olympia, Watershed Park, 15.iii.2018, em.?, E. Stansbury, ex Tolmiea menziesii, # CSE4607, CNC1135586 (1♀); Olympia, Watershed Park, 11.v.2018, em. 29.v.2018, ex Tellima grandiflora, E. Stansbury, # CSE5038, CNC934532–934534 (2♁ 1♀); Tolmie State Park, 23.v.2018, em. 8.vi.2018, E. Stansbury, ex Tiarella trifoliata, # CSE4610, CNC1144078 (1♁); same but em. 11.vi.2018, # CSE4609, CNC1135583 (1♁). Photographed mine. BRITISH COLUMBIA: Vancouver Island, Sooke, 10.viii.2019, Mews, Tiarella trifoliata, iNat 30890584. Hosts. Saxifragaceae: Heuchera micrantha Douglas, Tellima grandiflora (Pursh) Douglas ex Lindl. (Spencer 1981), Tiarella trifoliata L., Tolmiea menziesii (Pursh) Torr. & A.Gray (Griffiths 1972). Leaf mine. Upper-surface; entirely linear, “appearing white in incident light, up to 20-25 cm long, about 2 mm wide terminally; faeces scattered as discrete particles (mostly separated by over 1 mm), or forming short “threads” (Fadenstucke) in terminal part of mine” (Griffiths 1972). Puparium. Brown or white, with darker strip on ventral surface; formed within the leaf, on the lower surface, with its anterior spiracles projecting ventrally through the lower epidermis (Griffiths 1972). Phenology and voltinism. In Alaska, larvae and puparia collected in late August yielded one adult in early September and one the following spring. From this, Griffiths (1972) concluded that this species is at least partly multivoltine. The Washington material collected in March likely involved puparia that had overwintered, with the next generation represented by the mines collected in late May, which yielded adults in early to mid-June. Distribution. USA: AK (Griffiths 1972), CA (Spencer 1981), [OR] (see below), WA; [Canada: BC]. Comments. This species was reported from Washington previously, but based only on a female specimen (Winkler et al. 2009). The California records of Spencer (1981) are based on a male and female reared from Heuchera micrantha in Santa Cruz Co., along with leaf mines found on Tellima grandiflora in Marin Co. Griffiths (1972) also reported the latter host based only on leaf mines, and the CSE5038 specimens are the first actually to be reared from Tellima. Eiseman & Lonsdale (2018) reported as “ Phytomyza sp. 7” a female reared from Tiarella trifoliata in Oregon that could not be confidently identified as P. tiarellae, but this specimen does fit within the known range of P. tiarellae and we thus have not seen evidence that there is another species west of the Rocky Mountains whose leaf mine could be mistaken for that of P. tiarellae., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 38-39, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Spencer, K. A. (1981) A revisionary study of the leaf-mining flies (Agromyzidae) of California. University of California Division of Agricultural Sciences Special Publication, 3273, 1 - 489.","Griffiths, G. C. D. (1972) Studies on boreal Agromyzidae (Diptera). I. Phytomyza miners on Saxifragaceae. Quaestiones entomologicae, 8, 67 - 80.","Winkler, I. S., Scheffer, S. J. & Mitter, C. (2009) Molecular phylogeny and systematics of leaf-mining flies (Diptera: Agromyzidae): delimitation of Phytomyza Fallen sensu lato and included species groups, with new insights on morphological and host-use evolution. Systematic Entomology, 34, 260 - 292. [including online Supporting Information] https: // doi. org / 10.1111 / j. 1365 - 3113.2008.00462. x","Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1"]}

Ophiomyia nabali Eiseman & Lonsdale, spec. nov. (Figs. 24–28, 122–126) Holotype. USA. IOWA: Winneshiek Co., Decorah, Twin Springs Park, 24.viii.2017, em.?, J. van der Linden, ex Prenanthes alba, # CSE4667, CNC1135652 (1♁). Paratype. IOWA: Winneshiek Co., Decorah, Twin Springs Park, 19.x.2017, em. spring 2018, J. van der Linden, ex Prenanthes alba, # CSE4641, CNC1144031 (1♁). Other material examined. IOWA: Winneshiek Co., Decorah, Twin Springs Park, 10& 13.ix.2017, em. spring 2018, J. van der Linden, ex Prenanthes alba, # CSE4668, CNC1144030 (1 empty puparium), same data as paratype, # CSE4641, CNC1144032 (1 partial puparium). Etymology. The specific epithet refers to the host plant genus, Nabalus Cass. Host. Asteraceae: Nabalus albus (L.) Hook. Larval biology. The larva mines in the stem (CSE4641) or bores in the petiole (CSE4667, CSE4668). In the latter case, slight discoloration is evident externally, and in one instance (which first alerted us to this species’ exist- ence), the petiole was partially severed and scabbed over. One petiole gallery was observed to begin as a narrow mine extending ~ 2.5 cm into the leaf midrib. After opening the petiole to inspect this larva, it was successfully transferred to the midrib of a dandelion leaf (Taraxacum officinale F.H.Wigg.), where it fed for over a week before perishing. At one point its linear mine wandered into the leaf blade and then back into the midrib, but this species has otherwise been observed only to feed in the midrib, petiole, and stem of Nabalus. Puparium. (Figs. 27–28) Black; formed just beneath the petiole or stem epidermis, through which the anterior spiracles protrude. Phenology and voltinism. Larvae have been found beginning in late August. Pupae overwinter, with adults emerging in spring. Distribution. USA: IA. Adult description. Female unknown, wings of male not expanded; specimens died soon after emergence and are undeveloped, with one in poor condition; one specimen represented by partial puparium in preserved host stem; costa presumably extending to M 1, wing length likely to be approximately 2.3–2.7 mm based on body size. Eye height divided by gena height: 5.3. First flagellomere small, rounded. Ocellar triangle reaching level of posterior ori. Face with pronounced ridge with medial bulb; bulb narrow, with very thin medial groove that continues onto lunule where it is bordered by shallow ridges. Gena produced, forming nearly 60° angle; apex narrower and slightly pronounced. Cheek developed. Clypeus very narrow and apically truncated with arms bowed. Thorax subshining. Chaetotaxy: Two ori, two ors. Fasciculus developed, strongly upcurved on apical ¼ where hairs separate and appear frayed. Ocellar and postvertical setae subequal to ors. Two strong dorsocentral setae. Approximately ten rows of acrostichal setulae. Mid tibia with one small posteromedial seta. Coloration: (Figs. 24–26) Setae dark brown. Body dark brown, including halter. Calypter margin and hairs dark brown. Genitalia: (Figs. 122–126) Surstylus short, rounded, fused to distoventral margin of epandrium, with several irregular rows of tubercle-like setae on inner-distal surface. Hypandrium with long, laterally bulging arms; distal region subtriangular with pointed apex; inner lobe twisted, U-shaped with several sockets. Phallophorus narrow with swollen venter and strongly constricted base. Basiphallus mostly consisting of strong right-lateral band that reaches level of distiphallus; very short, weak, ill-defined left lateral band present; base truncated along margin of phallophorus. Mesophallus small, cylindrical, fused to ventromedial surface of distiphallus. Distal half of distiphallus relatively long and well-sclerotized with lateral texturing, narrowing apically to broadly rounded apex that is slightly upcurved; with thick ventral ridge along midline; with slightly swollen medial chamber at level of mesophallus that extends posterodorsally and is flared behind dorsal opening; widest at base, which is mostly clear except for scattering of minute sclerotized patches that concentrate ventrally. Ejaculatory apodeme not found. Comments. We have found similar Ophiomyia mines and puparia in stems and peduncles of Nabalus trifoliolatus Cass. in MA, NH, and NY, possibly representing O. nabali. The only other Ophiomyia known from Nabalus is O. congregata (Malloch), which has a fringe of peristomal hairs in the male and no fasciculus. Whereas O. nabali feeds exclusively in the stem, petiole, and leaf midrib, overwintering as a black puparium, O. congregata feeds partly as a leafminer, overwintering as an immature larva in the crown of the plant and forming a whitish puparium in the spring (Eiseman & Lonsdale 2018). The unfortunate state of preservation of known specimens of Ophiomyia nabali prevents the confident use of existing keys for identification, but the genal angle (60°), presumed wing length and groove along the facial keel key the holotype to O. lacertosa Spencer using Spencer & Steyskal (1986). The strongly bent fasciculus and unusual phallic morphology would eliminate this as a possibility, as they exclude the possibility of including the type in any other known species. The mesophallus is somewhat small and inserted far forward on the distiphallus, but it is the distiphallus itself that is most characteristic., Published as part of Eiseman, Charles S., Lonsdale, Owen, Linden, John Van Der, Feldman, Tracy S. & Palmer, Michael W., 2021, Thirteen new species of Agromyzidae (Diptera) from the United States, with new host and distribution records for 32 additional species, pp. 1-68 in Zootaxa 4931 (1) on pages 18-19, DOI: 10.11646/zootaxa.4931.1.1, http://zenodo.org/record/4545337, {"references":["Eiseman, C. S. & Lonsdale, O. (2018) New state and host records for Agromyzidae (Diptera) in the United States, with the description of thirty new species. Zootaxa, 4479 (1), 1 - 156. https: // doi. org / 10.11646 / zootaxa. 4479.1.1","Spencer, K. A. & Steyskal, G. C. (1986) Manual of the Agromyzidae (Diptera) of the United States. USDA Agricultural Research Service Agriculture Handbook, 638, 1 - 478. https: // doi. org / 10.5962 / bhl. title. 119606"]}