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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"]}

Zavrelia parapentatoma sp. nov. Figs. 3–4 LSID: urn:lsid:zoobank.org:act: 3AEFFA97-108E-493C-91A4-7CCCE57A5E91 Type material. Holotype male; USA, Michigan, Washtenaw Co., Ervin-Stucki Preserve; 42.136730°, -83.981339°; 8.vii.2021; leg. Patrick Hudson; deposited at ARC. Paratypes: 3 males, 1 female; same as holotype. 1 male, 1 female; USA, Michigan, Washtenaw Co., Ervin-Stucki Preserve; 42.136730°, -83.981339°; 8.v.2022; leg. Patrick Hudson; deposited at ARC. 5 pupae, 5 larvae; USA, Michigan, Washtenaw Co., Ervin-Stucki Preserve; 42.136730°, -83.981339°; 8.vi.2022; leg. Patrick Hudson; deposited at ARC. Diagnostic characters. Zavrelia parapentatoma can be separated from other Zavrelia species by the following combination of characters: Adult male with wing length ~ 1.3 mm, 4–5 longer than broad; AR 1.19; frontal tubercle minute, ~4 μm long; anal point with numerous (35) long spinulae (at least 8 times as long as wide) placed between long anal crest and flexing with their pointed ends directed anteriorly; 8 long median tergite setae placed on 1–3 light roundish fields at mid tergite IX; median volsella short, stout, with simple and subulate lamellae. Adult female with AR 0.23–0.28; temporal setae 8; frontal tubercle small, ~6 μm long; sternite VIII with 30 setae; notum including rami 131–170, 151 μm long; coxosternapodeme with developed anterior and posterior lobes. Pupa with cephalic tubercle weak and short; the median patch of microspinules much more prominent on tergites II–V but not wellextended to the lateral edge; those in II-IV appear divided with only a few microspinules in between; tergite II with small anterior patches of lateral shagreens; lateral shagreens prominent and extend along the lateral edge of tergites III–VI; segment VIII with a simple or bifid posterolateral spur; anal lobe seta taeniate. Larva with AR 0.93–1.23; AAR 0.64–0.77, 0.70; AHR 0.31–0.36, 0.30; ventromental plates medially reaching the border of the second and third lateral teeth of mentum; MVR 1.00–1.18, 1.11. Etymology. The new species is named after its similarity to Z. pentatoma. The prefix “ para ” is Latin, meaning “near” or “close”. Description. Male (n = 5). Total length 1.7–1.9, 1.8 mm. Wing 1.3 mm long and 0. 3 mm wide. Coloration of the mounted specimen. Head, thorax, and tergites brown. Legs and sternites yellowish brown. Wing and halters greyish-brown. Head. Plume well-developed; antenna with 10 flagellomeres, ultimate flagellomere with 2 sensilla clavata (Fig. 3A); AR 1.12–1.29, 1.19. Frontal tubercle minute, 4 μm long. Temporal setae with only 2 inner verticals. Tentorium 99–110, 105 μm long (Fig. 3B). Clypeus squared about 82 μm long and 89 μm wide, with 10 setae 78–94 μm long. Third palpomere with 1 sensillum clavatum; lengths of palpomeres (in μm): 46–70, 53; 30–42, 38; 93–116, 105; 76–88, 83; 92–106, 97. Thorax chaetotaxy. Ac 12; Dc 9; Pa 1; Scts 4–5. Wing (Fig. 3C). Wing 4–5 times longer than wide. R with 17–20 setae, R 1 with 6–7, R 4+5 with 18–19 setae. VR 1.5. Wing cell setation mainly confined to the wing apex. Legs. Fore leg tibia with 40 μm long spur; mid and hind legs tibiae with well-separated combs; mid leg tibial combs 12 μm long, and spurs 13–16 μm long; hind leg tibia with combs 9–11 μm long and spurs 13–17 μm long. Lengths and ratios of leg segments as in Table 1. Hypopygium (Fig. 3D–E). Tergite IX 103–129, 114 μm long and 60–86 μm wide, with 8 median setae on 1–3 light roundish fields, and 20 apical setae. Anal point 42–58, 50 μm long and 20–24, 22 μm wide (maximum); with 2 lateral setae close to the base; well-developed crests present; 33–37, 35 very long spinulae present between crests of anal point (Fig. 3D–E). Gonocoxite 64–70, 67 μm long; gonostylus 47–54, 49 μm long; HR 1.30–1.5, 1.4. Superior volsella 42–45 μm long, digitiform, medially directed, with 3 dorsal and 3 median setae on setiger; median volsella stout with cluster of simple and subulate, lamella, stem 16–19, 17 μm long; inferior volsella club-shaped, 37–52, 49 μm long with several distal setae, dorsal surface without microtrichia. HV 3.6. Female (n = 2). Total length 1.8–1.9 mm. Wing 1.3 mm long and 0.32 mm wide. The coloration of the mounted specimen. Same as male. Head. Antennae with 5 flagellomere (Fig. 3F) with ultimate flagellomere 59–61 μm long, AR 0.23–0.28, 0.26. Temporal setae 8. Frontal tubercle small, 6 μm long. Tentorium 107–116, 112 μm long. Clypeus with 12 setae, setae 77 μm long. Lengths of palpomeres (in μm): 40–49, 44; 27–37, 32; 71–80, 76; 62–63; 94. Thorax chaetotaxy. Ac 8; Dc 7; Pa 1; Scts 6. Wing. Wing L/ W 3.8 –4.0. Brachiolum with 2 setae; R with 19 setae, R4+5 with 24 setae, R1 with 7 setae. Wing membrane setation same as in male. VR 1.5–1.6. Legs. Foreleg tibial spur with 23 μm long; mid and hind legs tibiae with well-separated combs; mid leg tibial spurs 16 μm long; hind leg tibial spurs 9–11 μm long. Lengths and ratios of leg segments as in Table 2. Genitalia (Fig. 3G). Tergite IX 138–163, 151 μm long. Sternite VIII with 30 setae. Gonocoxite with 1 seta. Notum including rami 131–170, 151 μm long, notum alone 83–102, 92 μm long. Seminal capsule 56 μm in diameter (n = 1). Coxosternapodeme with developed anterior and posterior lobes. Cercus 49 μm long and 30 μm wide. Pupa (n = 5). Total length 2.2–2.3 mm; abdomen 1.6–1.7 mm long. Colouration of the pupal exuviae. Pale brown. Lateral margins of abdominal segment VIII much darker. Cephalothorax: Cephalic tubercle weak and short, 18–29, 22 μm. Taeniate frontal seta 135–149, 142 μm long (Fig. 4A). Pedicel sheath tubercle absent. Thoracic horn 248–428, 349 μm long, 18–28, 22 μm wide, with few spines in mid-section (Fig. 4B); precorneals taeniate, Pc 1–3 136, 183, 196 μm long (n = 1). Wing sheath 699–754, 725 μm long and 177–195, 187 μm wide. Abdomen (Fig. 4C): Tergite I bare. Tergites II–IX with median patch of microspinules. Median patch of microspinules much more prominent on tergites II–V and not well extended to the lateral edge; those in II–IV appear divided in the middle with only a few microspinules; the median patch of microspinules on tergites VI–IX reducing gradually and well-divided. Pleura II with small anterior patches of lateral shagreens; lateral shagreens prominent and extend along the lateral edge of pleura III–VI. No visible lateral shagreens on pleura VII–IX. Tergite II with pedes spuri B and posterior hook row of 50–60, 55 hooks, 120–155, 137 μm wide. Sternites bare. Segment II–III with 3 simple lateral setae; segment IV with 2 simple and 1 taeniate lateral setae; segment V–VII with 4 taeniate lateral setae; segment VIII with 3 taeniate lateral setae and a simple to bifid posterolateral spur, spur 29–35, 32 μm long. Anal lobe 108–123, 116 μm long and 61–83, 74 μm wide; genital sac 163–190 μm long and 50–69 μm wide; anal lobes with fring of 16–17 taeniate setae of 73–112, 90 μm long; anal lobe seta taeniate 77–81, 79 μm long. Larva (n = 5). Total length 2.9–3.3, 3.0 mm, case 2.9 mm long. Coloration. Head capsule brown, mentum, inner teeth of mandibles, and occipital region darker. Abdomen yellowish brown. Head: HL/HW 0.84–1.05, 0.9. AR 0.93–1.23, 1.1; antennal pedestal 55–64, 57 μm long with well developed, 29–31, 30 μm long spur; antennal segment lengths (in μm): 73–88, 82; 32–40, 37; 18–22, 19; 11–15, 13; 7–8; third segment inserted subapically on segment two; AAR 0.64–0.77, 0.70; AHR 0.31–0.36, 0.30; antennal blade 113–114 μm long; Lauterborn organs 17–26, 22 μm long (Fig. 4D). Labral SI comb-like, SII plumose, SIII simple. S3 simple, 86–104, 95 μm long. Premandible 49–53, 51 μm long; quartered and with a well-developed brush (Fig. 4E). Mandible 82–88, 84 μm long, with 1 subapical, 1 apical, and 3 inner teeth; setae subdentalis 48–56, 51 μm long, reaching well-beyond apical tooth; setae interna with 3 blades of well-branched spines (Fig. 4F). Mentum with one median tooth and six lateral teeth; first lateral teeth slightly lower than second and third; ventromental plates medially reaching the border of the second and third lateral teeth of mentum (Fig. 4G); MVR 1.00–1.18, 1.11. Body: Posterior parapod 86–102, 94 μm long and 124–133, 130 μm wide, with 16 simple claws. Procercus 23–29, 24 μm long and 23–29, 26 μm wide; each procercus with 4 apical setae, 2 shorter 372–404, 380 μm long, and the 2 longer 502–679, 617 μm long; subapical seta 163–188, 171 μm long; supranal seta 237–255, 248 μm long. 4 conical anal tubules present, 73–92, 82 μm long. Taxonomic remarks. The Z.parapentatoma and Z.pentatoma are close species and probably form a sister group. The morphological characters that separate the two species are, at the most, very subtle. There are many overlapping sizes and ratios between both species, which makes the distinction between the two species difficult (Table 3). For adult males, the only distinguishing character is the spinulae of the anal point, which in Z. parapentatoma are much longer (at least 8 times as long as wide) compared to Z. pentatoma (no more than 4 times as long as wide). Females of both species are hard to separate; the range of notum and ramus length combined may be slightly different between the two species, with Z. parapentatoma being in the higher range. The wing venarum ratio of adults of both species also differs slightly, with Z. parapentatoma having the higher ratio. The pupae of both species show the most significant difference among life stages. Z. parapentatoma has median patches of microspinules on tergites II–V less extended, and those in II–IV appear divided in the middle with only a few microspinules. Additionally, the anal lobe seta is prominent and taeniate in Z. parapentatoma. Z. pentatoma has a well-extended and joined patch of microspinules on tergites II–V, and the anal lobe seta is simple. Few characters could separate the larva of Z. parapentatoma from that of Z. pentatoma including the extent of the ventromental plates, size of Lauterborn organs, length of the shortest anal setae, and length of the anal tubules. In Z. parapentatoma ventromental plates medially reach the border of the second and third lateral teeth of the mentum, whereas in Z. pentatoma hardly reach the third lateral teeth. Z. parapentatoma has a slightly longer and narrower Lauterborn organ, and its shortest-anal setae and anal tubules are also longer than those of Z. pentatoma., Published as part of Namayandeh, Armin, Hudson, Patrick L., Ghaderi, Edris & Stott, Wendylee, 2023, Zavrelia parapentatoma (Chironomidae: Diptera), a curious new species from North America, revealed by molecular methods, pp. 111-124 in Zootaxa 5249 (1) on pages 115-118, DOI: 10.11646/zootaxa.5249.1.6, http://zenodo.org/record/7685359

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"]}

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."]}

Key to adult males of Zavrelia 1. Anal point densely covered with strong spinulae............................................................. 2 - Anal point bare, with microtrichia or short spinules only...................................................... 3 2. Anal point spinulae of varying size and shapes but never very long, ≤ 4 × as long as wide (Ekrem and Stur, 2009: Fig. 8E; Giłka 2008: Fig. 4)........................................................ Z. pentatoma Kieffer & Bause (Palearctic) * - Anal point spinulae very long, ≥ 8 × as long as wide (Fig. 3D–E).................... Z. parapentatoma sp. n. (Nearctic) * 3. Setiger of superior volsella with conspicuous constriction in apical 1/3........................................... 4 - Setiger of superior volsella without conspicuous constriction in apical 1/3........................................ 6 4. Setiger of superior volsella with pointed apex (Ekrem and Stur, 2009: Fig. 5E)...... Z. clinovolsella Guo & Wang (Oriental) - Setiger of superior volsella with somewhat rectangular apex................................................... 5 5. Anal point with only a few microtrichia in between crests; distinct microtrichia-free areas on anal tergite around the base of anal point (Ekrem and Stur, 2009: Fig. 10E)......................................... Z. sinica Ekrem & Stur (Palearctic) - Anal point with numerous microtrichia in between crests; microtrichia present all around the base of anal point (Zorina 2008: Figs. 13–14).......................................................... Z. pseudopentatoma Zorina (Palearctic) 6. Anal point with small spinules in between crests............................................................. 7 - Anal point bare, or with microtrichia in between crests at most................................................. 9 7. AR close to 0.6................................................................ Zavrelia casasi Ekrem & Stur - AR> 0.7............................................................................................ 8 8. AR 1.23; wing length about 1.50 mm; LR 1 about1.45; laterosternite with one seta; anal point with small spinules scattered between the entire length of anal crests (Ekrem & Stur 2009: Fig. 12E)..... Z. tusimatijea (Sasa & Suzuki, 1999) (Palearctic) - AR 0.91–1.08; wing length 0.88–1.04 mm; LR1 about 2.09; laterosternite without seta; anal point with small spinules scattered between anal crests at anal point base only (Guo and Wang, 2007: Fig. 3; Lin & Wang 2017: Figs. 8–9)........................................................................................ Z. bragremia Guo & Wang (Oriental) 9. Anal point bare...................................................................................... 10 - Anal point with microtrichia in between crests............................................................. 11 10. AR 1.00–1.18; LR 1 1.36–1.46; superior volsella with pointed apex (Zorina, 2008: Figs. 1–2).... Z. elenae Zorina (Palearctic) - AR 0.45; LR1 1.96; superior volsella with rounded apex (Kobayashi 2014: Fig. 7)......................................................................................... Z. simantoneoa (Sasa, Suzuki & Sakai, 1998) (Palearctic) 11. Wing length 1.40 mm; AR close to 0.75....................................... Z. hudsoni Ekrem & Stur (Nearctic) - Wing length close to 1.00 mm; AR close to 0.90................................. Z. aristata Ekrem & Stur (Nearctic), Published as part of Namayandeh, Armin, Hudson, Patrick L., Ghaderi, Edris & Stott, Wendylee, 2023, Zavrelia parapentatoma (Chironomidae: Diptera), a curious new species from North America, revealed by molecular methods, pp. 111-124 in Zootaxa 5249 (1) on page 120, DOI: 10.11646/zootaxa.5249.1.6, http://zenodo.org/record/7685359, {"references":["Ekrem, T. & Stur, E. (2009) A review of the genus Zavrelia (Diptera: Chironomidae). European Journal of Entomology, 106, 119. https: // doi. org / 10.14411 / eje. 2009.016","Gilka, W. (2008) An intraspecific morphological variability of Zavrelia pentatoma Kieffer (Diptera: Chironomidae). Dipteron, Bulletin of the Dipterological Section of the Polish Entomological Society, 24, 11 - 15.","Zorina, O. V. (2008) Russian Zavrelia Kieffer (Diptera: Chironomidae), with the description of two new species. Zootaxa, 1845 (1), 60 - 68. https: // doi. org / 10.11646 / zootaxa. 1845.1.4","Lin, X. L. & Wang, X. H. (2017) A redescription of Zavrelia bragremia Guo & Wang, 2007 (Diptera: Chironomidae). CHIRONOMUS Journal of Chironomidae Research, 30, 67 - 71. https: // doi. org / 10.5324 / cjcr. v 0 i 30.2345","Kobayashi, T. (2014) A redescription of Zavrelia simantoneoa (Sasa, Suzuki and Sakai, 1998) comb. nov. CHIRONOMUS Journal of Chironomidae Research, 27, 41 - 44. h"]}

Genus Cedrimyia gen. nov. Figs. 1���2 LSID: urn:lsid:zoobank.org:act: 7845E8DA-9E57-427D-80C5-BEE7B922A672 Type species. Cedrimyia margareti Namayandeh and Hudson, sp. n. by present designation. Diagnostic characters. A combination of eyes with wedge-shaped dorsomedial extension; presence of 2 scalpellate acrostichals in centre of scutum; anterpronotal lobes without gap; tergite IX with curved ridge running close to mid-section of anal point; anal point long, bearing simple, short and weak setae, pointing posteriorly; virga consisting of two long spines; gonostylus with strong, comparatively darker spine-like setae; crista dorsalis thin and blade-like, stretching from mid to anterior of gonostylus separates Cedrimyia from most Orthocladiinae (see also the remarks section). Etymology. The new genus is named after its type locality, Cedar Falls in Ohio. The suffix ��� myia ��� is Greek for the fly. Description. Adult male. Small to medium size species up to 3.2 mm long. Wings up to 2.3 mm long. Head (Fig. 1a). Antenna with 12 flagellomeres, 2 nd ���3 rd segments and last flagellomere with sensilla chaetica. Eyes bare, with wedge-shaped dorsomedial extension. Temporal setae present, including inner verticals, postoculars and outer verticals. Tentorium narrow with small tentorial pit close to apex. Clypeus rectangular. Palp 5 segmented with third palpomere having sensilla clavata. Thorax (Fig. 1b). 2 scalpellate acrostichals in centre of scutum. Dorsocentrals in single row, prealars few, scutellars in single row. Humeral pit small. Antepronotal lobes developed, narrowing anteriorly, without gap and bare. Wing (Fig. 1c). Brachiolum, squama and R with setae. Other veins without setae. Costa extended. R 4+5 ends just above M 3+4. Anal lobe squared. Microtrichia visible at> 100 x magnification. Legs. Mid and hind tibia with long sparse beard, hind femur with keel. Pulvilli absent. Fore tibia with 1 spur, each of mid and hind tibia with 2 spurs. Hind tibia comb developed. Hypopygium (Fig. 1d). Tergite IX with curved ridge running close to mid-section of anal point. Anal point narrowly triangular, long, reaching close to the apex of inferior volsella, bearing simple, short, and weak setae, distributed evenly along its mid-section, and pointing posteriorly, apex rounded. Virga present consists of two long spines. Sternapodeme straight and thick. Phallapodeme collar bone-shaped. Inferior volsella a prominent triangular lobe. Gonostylus squat-shaped, well expanded in mid to anterior section, surface with strong, comparatively darker, spine-like setae; crista dorsalis developed, thin and blade-like, stretching from mid to anterior of gonostylus. Adult female. Small species up to 2.1 mm long. Wing up to 2.4 mm long. Head (Fig. 2 a-b). Antenna with 5 flagellomeres, 3 rd ���4 th segments and last flagellomere with sensilla chaetica. Eyes bare, with short dorsomedial extension. Temporal setae present, including inner verticals, postoculars, orbitals and outer verticals. Tentorium long and narrow with small tentorial pit close to apex. Clypeus rectangular. Palp 5 segmented with third segment bearing sensilla clavata. Thorax. 2 scalpellate acrostichals in centre of scutum. Dorsocentrals in single row, prealars few, scutellars in single row. Humeral pit small. Antepronotal lobes developed, narrowing anteriorly, without gap and bare. Wing (Fig. 2c). Brachiolum, squama, R and R 4+5 with setae. Other veins without setae. Costa extended. R 4+5 ends just above M 3+4. Anal lobe squared. Microtrichia visible at> 100 x magnification. Legs. Mid and hind tibia with long sparse beard, hind femur with keel. Pulvilli absent. Fore tibia with 1 spur, each of mid and hind tibia with 2 spurs. Hind tibia comb developed. Genitalia (Fig. 2 d-e). Gonocoxite narrow and long with few setae. Seminal capsules small, semi-circular, spermathecal ducts without loops. Gonapophysis VIII divided into large ventrolateral lobe covering the base of smaller dorsomesal lobe. Apodeme lobe distinct. Tergite IX divided in 2 crescent-shaped setigerous light patches. Cercus with circular base and pediform extensions. Remarks. The short and weak anal point setae; long, thin, and blade-like crista dorsalis of gonostylus; and the presence of dorsomedial extension of the eyes are probably autapomorphic in Cedrimyia. The presence of strong, comparatively darker setae on the surface of the Cedrimyia gonostylus can also be considered autapomorphic. The presence of scalpellate acrostichals on scutum and long anal point of adult males resembles the species in Antillocladius Saether, 1981, Litocladius Mendes, Andersen & Saether, 2004, Lyrocladius Mendes and Andersen, 200, and Compterosmittia Saether, 1981. The adult male of Cedrimyia can be distinguished from Antillocladius and Litocladius by a combination of short dorsomedial extension of the eye; only 2 scalpellate acrostichals in centre of scutum; short, weak, and posteriorly directed setae of anal point; virga consisting of two long and prominent spines; and thin blade-like crista-dorsalis of the gonostylus. The absence of lateral lamella of virga can further separate adult males of Cedrimyia from Litocladius. The setose squama of Cedrimyia adults can further separate them from Compterosmittia (See Mendes et al. 2004). The presence of strong, comparatively darker setae on the surface of the Cedrimyia gonostylus is also a distinguishing character. Species of Lyrocladius have a laterally located strong setae on their gonostylus. The female of Cedrimyia can be distinguished from Antillocladius by a divided tergite IX with two setigerous protrusions, and a more prominent dorsomesal lobe of gonapophysis IX separates it from Litocladius. In their review of the Orthocladiinae genera with scalpellate acrostichals Mendes and Andersen (2008) placed Antillocladius, Litocladius, and Lyrocladius Mendes and Andersen, 2008 in the same monophyletic group based on the apomorphic characters such as scalpellate acrostichals, and similarity in the shape of anal point and its setation. Cedrimyia likely forms a sister group to this group. Among other Orthocladiinae, the long anal point with weak lateral setae and the wedge-shaped eye extension bears some resemblance to Paraphaenocladius Thienemann, 1924. A longitudinal ridge on tergite IX of adult males may resemble the species in Tavastia Tuiskunen, 1985, and Mesosmittia Brundin, 1956. Bare eye, absence of apical setae on the last antennal flagellomere, and bare wing of the adults separate Cedrimyia from those of Tavastia. The presence of scalpellate acrostichals in centre of scutum and a more prominent anal point separates Cedrimyia from those of Mesosmittia., Published as part of Namayandeh, Armin & Hudson, Patrick L., 2022, Two new Orthocladiinae (Diptera: Chironomidae) collected in overlooked places of Midwest USA, pp. 120-128 in Zootaxa 5099 (1) on pages 121-123, DOI: 10.11646/zootaxa.5099.1.5, http://zenodo.org/record/6325512, {"references":["Mendes, H. F., Andersen, T. & Saether, O. A. (2004) A review of Antillocladius Saether, 1981; Compterosmittia Saether, 1981 and Litocladius new genus (Chironomidae, Orthocladiinae). Zootaxa, 594, 1 - 82. https: // doi. org / 10.11646 / zootaxa. 594.1.1","Mendes, H. F. & Andersen, T. (2008) A review of Antillocladius Saether and Litocladius Mendes, Andersen et Saether, with the description of two new Neotropical genera (Diptera, Chironomidae, Orthocladiinae). Zootaxa, 1887 (1), 1 - 75. https: // doi. org / 10.11646 / zootaxa. 1887.1.1"]}

Parakiefferiella mishigami sp. nov. Fig. 3 LSID: urn:lsid:zoobank.org:act: 33D7B56A-63BF-4F0E-88B5-DDD24DC2C870 Type material. Holotype ♂; 19.V.1991; USA, Michigan, Chippawa Co., Neebish Island, St. Marys River, Downbound channel; 46.292, -84.218; leg. Patrick Hudson; deposited at ARC. Diagnostic characters. The new species can be separated from other Parakiefferiella species by a combination of the following characters: AR 1.1; costa of wing not well-extended; LR 1 0.42; anal point shallow, broadly triangular; virga with two short spines; inferior volsella bilobed with dorsal lobe smaller, anterior to a larger, roundly projecting ventral lobe. Etymology. The new species is named after the state of Michigan. The name mishigami is Ojibwe, meaning a ���large lake���. Description. Male (n=1). Total length 2.9 mm. Wing 2.1 mm long and 0.51 mm wide. Coloration of mounted specimen. Head, thorax, halter and tergites brown. Legs and sternites light brown. Wing greyish-brown Head (Fig. 3a). Antenna with 13 flagellomere, last flagellomere with 12 sensilla chaetica, 2 nd ���3 rd segments each with 2 sensilla chaetica, groove starts at 4 th segment, AR 1.1. Eyes bare, without dorsomedial extension, temporal setae with 4 inner verticals in single row (Fig. 3a). Tentorium narrow with large tentorial pit close to apex (Fig. 3a), tentorium 141 ��m long. Clypeus rectangular, 71 ��m long and 123 ��m wide, bearing 7 setae, setae 59���77, 67 ��m long. Palpal segments lengths (in ��m): 56, 37, 80, 83, 117. Thorax. Dorsocentrals 5, prealars 4, scutellars 6 in single row. Median tuft of setae in centre of scutum (Fig. 3b). Humeral pit vestigial. Antepronotal lobes developed, narrowing anteriorly, with small gap and bearing 3 setae. Wing. Brachiolum bare. Squama bare. R and R 1 apparently bare, other veins without setae. Costa not well-extended. Anal lobe rounded. Fine punctation visible at 40 x magnification. Legs. Femur and tibia of all legs with long beard. Pulvilli vestigial. Fore tibia spur 40 ��m long, mid tibia spurs 11 ��m long, hind tibia spurs 38 and 15 ��m long, hind tibia comb with around 18 spines. Lengths and proportions of legs as in Table 3. Hypopygium (Fig. 3c). Anal point shallow, broadly triangular, bearing around 10 simple setae, anal point 20 ��m long and 46 ��m wide. Virga consists of two spines, 21 ��m long. Sternapodeme slightly arched with large oral projections, sternapodeme 98 ��m long. Phallapodeme large, 51 ��m long. Inferior volsella consist of two lobes, the dorsal lobe smaller, digitiform, located anteriad to the larger, roundly projecting ventral lobe. Gonostylus bent, tapered posteriorly, gradually expanding anteriorly, gonostylus 82 ��m long, crista dorsalis receded. Gonocoxite 193 ��m long. HR 2.3, HV 3.4. Remarks. Combination of a tuft of setae in centre of scutum, eyes bare and without dorsomedial extension, a bare wing with bare squama, and vestigial pulvilli places this specimen in Parakiefferiella. This species can be separated from the species of Epoicocladius Zavřel, 1924 based on its relatively short apical palpal segment and a median bent in its gonostylus. The presence of a well-separated bilobed inferior volsella can be seen in some species of Parakiefferiella, more prominently in Parakiefferiella bilobata Tuiskunen 1986. However, P. mishigami differs from these species in that the posterior lobe is much more prominent than the anterior lobe, and it also differs in shape. Ecology and habitat. The single adult collected in the vicinity of St. Marys River, Michigan, suggesting the larvae inhabiting large running waters., Published as part of Namayandeh, Armin & Hudson, Patrick L., 2022, Two new Orthocladiinae (Diptera: Chironomidae) collected in overlooked places of Midwest USA, pp. 120-128 in Zootaxa 5099 (1) on pages 125-127, DOI: 10.11646/zootaxa.5099.1.5, http://zenodo.org/record/6325512

Cedrimyia margareti sp. nov. Type material. Holotype ♂; 24.III.2010; USA, Ohio, Hocking Co., Hocking Hills State Park, Queer Creek below Cedar Falls; 39.4194, -82.5269; leg. Patrick Hudson; deposited at ARC. Paratype 3♂♂ same as the holotype. Paratype 1♂; 16.X.1990; USA, Wisconsin, Douglas Co., Superior Harbor, East End of Lake Superior; 46.691, -91.985; leg. Patrick Hudson; deposited at ARC. Paratype 2♀; 24.III.2010; USA, Ohio, Hocking Co., Hocking Hills State Park, Queer Creek below Cedar Falls; leg. Patrick Hudson; deposited at ARC. Etymology. The new species is named in honor of Margaret Hudson, who has been a great help in collecting type specimens, making this study possible. Description. Male (n = 5). Total length 2.8���3.2, 2.9 mm. Wing 2.3 mm long and 0.65 mm wide. Coloration of mounted specimen. Head, thorax, halter brown. Legs and abdomen light to golden brown. Wing light to golden-brown Head (Fig. 1a). Antenna with 12 flagellomeres, last flagellomere with 8 sensilla chaetica, 2 nd ���3 rd segments each with 2 sensilla chaetica, groove starts at 3 rd segment, AR 1.3���1.4. Eyes bare, with wedge-shaped dorsomedial extension. Temporal setae 9���10, including 3 inner verticals, 3���4 postoculars and 3���4 outer verticals. Tentorium narrow with small tentorial pit close to apex (Fig. 1a), tentorium 135���156, 146 ��m long. Clypeus rectangular, 87 ��m long and 116 ��m wide, bearing 5���7, 6 setae, setae 77���85, 80 ��m long. Palpal segments lengths (in ��m): 44���54, 50; 50���61, 57; 73���99, 81; 83���90, 85; 127���156, 148. Third palpomere with 1 sensilla clavata. Thorax (Fig. 1b). 2 scalpellate acrostichals in centre of scutum. Dorsocentrals 5���7, 6 in single row, prealars 2���3, scutellars 4 in single row. Humeral pit small. Antepronotal lobes developed, narrowing anteriorly, without gap and bare. Wing (Fig. 1c). Brachiolum with 2 setae. Squama with 5 setae. R with 5 and R 1 apparently bare, other veins without setae. Costa extended, extension 69 ��m. R 4+5 ends just above M 3+4. Anal lobe squared. Microtrichia visible at> 100 x magnification. Legs. Mid and hind tibia with long sparse beard, hind femur with keel. Pulvilli absent. Fore tibia spur 43���53, 47 ��m long, mid tibia spurs 24���25; 18���23, 20 ��m long, hind tibia spurs 44���51, 48 and 15 ��m long, hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 1. Hypopygium (Fig. 1d). Tergite IX with curved ridge running close to mid-section of anal point. Anal point narrowly triangular, long, reaching close to the apex of inferior volsella, bearing around 6 simple, short, and weak setae, distributed evenly along its mid-section, and pointing posteriorly, apex rounded; anal point 75���97, 84 ��m long and 27���32, 29 ��m wide. Virga present consists of two long spines, 50���54, 53 ��m long. Sternapodeme straight and thick, 65���73, 69 ��m long. Phallapodeme collar bone-shaped, 72���86, 81 ��m long. Inferior volsella a prominent triangular lobe. Gonostylus squat-shaped, well expanded in mid to anterior section, surface with 3���4 strong, comparatively darker, spine-like setae; crista dorsalis developed, thin and blade-like, stretching from mid to anterior of gonostylus; gonostylus 65���75, 69 ��m long. Gonocoxite 181���194, 188 ��m long. HR 3, HV 3. Female (n = 2). Total length 2.0��� 2.1 mm. Wing 2.1���2.4, 2.2 mm long and 0.76���0.86, 0.80 mm wide. Coloration of mounted specimen. Head, thorax, halter brown. Legs and abdomen light to golden brown. Wing light to golden-brown Head (Fig. 2 a-b). Antenna with 5 flagellomeres, last flagellomere with 10 sensilla chaetica, 3 rd ���4 th segments each with 2 sensilla chaetica (Fig. 2a), AR 0.5���0.6. Eyes bare, with short dorsomedial extension. Temporal setae 5, including 1 inner verticals, 1 postoculars, 2 orbitals and 1 outer verticals (Fig. 2b). Tentorium long and narrow with small tentorial pit close to apex, tentorium 126���149, 137 ��m long. Clypeus rectangular, 89 ��m long and 109���127, 118 ��m wide, bearing 5 setae, setae 88 ��m long. Palpal segments lengths (in ��m): 44���52, 48; 49���55, 52; 82���84, 83; 77���84, 80; 138���154, 146. Third palpomere with 1 sensilla clavata. Thorax. 2 scalpellate acrostichals in centre of scutum. Dorsocentrals 5���6 in single row, prealars 2, scutellars 4 in single row. Humeral pit small. Antepronotal lobes developed, narrowing anteriorly, without gap and bare. Wing (Fig. 2c). Brachiolum with 2 setae. Squama with 5 setae. R with 6���10, 8, R 1 with 2���3 setae, R 4+5 with 2���3 setae, other veins without setae. Costa extended, extension 137 ��m. R 4+5 ends just above M 3+4. Anal lobe squared. Microtrichia visible at> 100 x magnification. Legs. Mid and hind tibia with long sparse beard, hind femur with keel. Pulvilli absent. Fore tibia spur 22���30 (26) ��m long, mid tibia spurs 22���25, 23; 21���22 ��m long, hind tibia spurs 38���52, 45 and 22 ��m long, hind tibia comb with around 12 spines. Lengths and proportions of legs as in Table 2. Genitalia (Fig. 2 d-e). Gonocoxite narrow and long with 4 setae. Seminal capsules small, semi-circular, spermathecal ducts without loops (Fig. 2d), seminal capsule 35���46, 40 ��m long, 31 ��m wide. Notum 174 ��m long. Gonapophysis VIII divided into large ventrolateral lobe covering the base of smaller dorsomesal lobe. Apodeme lobe distinct. Tergite IX divided in 2 crescent-shaped setigerous light patches (Fig. 2e). Cercus with circular base and pediform extensions (Fig. 2e), 117 ��m long and 43���55, 49 ��m wide. Ecology and habitat. Adults of the species were collected in the vicinity of Queer Creek just below Cedar Falls and along Superior Harbor in Lake Superior, vastly different habitats. Queer Creek is a third-order stream running through a deep ravine with rugged sandstone cliffs covered in part with lichen and with numerous seeps. Superior Harbor is part of a highly modified St. Louis River estuary for maintaining a safe harbor for both Great Lakes and international shipping. Extrapolating from these suggests that the habitat for this new genus is probably not truly aquatic but is presumably decaying moist organic matter along the shores of these two habitats. It is likely that the high organic (humus) soil of this region provides an ideal habitat for terrestrial Chironomidae. In East Fork Queer Creek, Ohio, close to our sampling location, Bolton and Jacobsen (2010) previously found close to seven species of terrestrial Chironomidae. The presence of emerging adults in May and again in October indicates that this species has more than one voltine., Published as part of Namayandeh, Armin & Hudson, Patrick L., 2022, Two new Orthocladiinae (Diptera: Chironomidae) collected in overlooked places of Midwest USA, pp. 120-128 in Zootaxa 5099 (1) on pages 124-125, DOI: 10.11646/zootaxa.5099.1.5, http://zenodo.org/record/6325512, {"references":["Bolton, M. J. & Jacobsen, R. E. (2010) The immature stages of Chasmatonotus unimaculatus Loew (Diptera: Chironomidae: Orthocladiinae) with notes on their habitat. In: Ferrington Jr., L. C. (Ed.), Proceedings of the XV International Symposium on Chironomidae, St. Paul, Minnesota, 12 - 14 August 2003. University of Minnesota, St. Paul, Minnesota, pp. 255 - 261."]}

Namayandeh, Armin, Hudson, Patrick L. (2022): Two new Orthocladiinae (Diptera: Chironomidae) collected in overlooked places of Midwest USA. Zootaxa 5099 (1): 120-128, DOI: 10.11646/zootaxa.5099.1.5

Namayandeh, Armin, Beresford, David V. (2021): Review of the genus Doncricotopus (Orthocladiinae: Chironomidae): with the description of a new species from Canada. Journal of Natural History 55 (19-20): 1195-1211, DOI: 10.1080/00222933.2021.1939186, URL: http://dx.doi.org/10.1080/00222933.2021.1939186

Rheocricotopus (Rheocricotopus) effusus (Walker, 1856) Adults: Male described by Albu (1968), Lehman (1969), Saether (1985), Makarchenko and Makarchenko (2005), hypopygium in key by Pinder (1978). Female described by Saether (1985). Immatures: Pupa described by Saether (1985), in key by Langton (1991). Larva described by Cranston (1982), Saether (1985), and in key by Epler (2001). Ecology and Habitat: Larvae mainly inhabit springs (Thienemann 1954). NE: Canada (Alberta, Northwest Territories); USA (Illinois, North Carolina, South Carolina, South Dakota). Widespread in the Palearctic., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 19, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Albu, P. 1968. Chironomide din Carpatii romanesti (III). - Studii si Cercetari de Biologie, Serie de Zoologie. Bot. 20 (5): 455 - 465.","Pinder, L. C. V. 1978. A key to adult males of the British Chironomidae (Diptera), Vol. 1, The key; Vol. 2, Illustrations of the Hypopygia. - Freshwater Biological Association Scientific Publication 37: 1 - 169.","Langton, P. H. 1991. A key to pupal exuviae of West Palaearctic Chironomidae. Private publication. Printed by Graytones, UK. 386 p.","Cranston, P. S. 1982. A key to the larvae of the British Orthocladiinae (Chironomidae). - Scientific Publication Freshwater Biological Association 45: 1 - 152.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p."]}

Key to the known Nearctic adult male Parametriocnemus Goetghebuer Abbreviations: AnP = Anal Point; CD = Crista Dorsalis; Gc = Gonocoxite; Gs = Gonostyle; IVo = Inferior Volsella; SVo = Superior Volsella; T = Tergite. 1a. TIX with AnP short not reaching the IVo (Saether 1969, fig.62)... P. gramnicola (Lundbeck) 1b. TIX with AnP longer, reaching the IVo........... 2 2a. IVo narrow, finger-like with hooked apex (Fig. 1e; Sublette 1967, fig. 35)................................................................................ P. hamatus (Johannsen) 2b. IVo with broad base and rounded apex............ 3 3a. AnP very long reaching beyond IVo; CD enlarged, broadly projecting above the inner margin of Gs (Saether 1969, fig. 65).................................................................................. P. vespertinus Saether 3b. AnP short, reaching only as far as IVo; CD either inconspicuous or present as small preapical to oth......................................................................... 4 4a. IVo with a broad basal attachment about half the length of Gc. AnP with weak lateral setae (Saether 1969, fig. 63; Sublette 1967, fig. 33).......................................... P. lundbeckii (Johannsen) * 4b. IVo with narrow basal attachment about quarter length of Gc. AnP with bristle-like setae.......... 5 5a. Gs angular distally with dorsal ridge ending in small sharply pointed CD (Saether 1969, fig. 58)..................................................... P. eoclivus Saether 5b. Gs somewhat rounded distally with weak dorsal margin merging with poorly defined CD (Gowin and Thienemann 1942, fig. 2)........................................... P. boreoalpinus Gowin and Thienemann *Will also key to Parametriocnemus stylatus (Spärck, 1923), a Palearctic species. Parametriocnemus lundbeckii and P. stylatus closely resemble each other. The two species can partially be separated based on the characters of gonostyle. The gonostyle of P. stylatus is much narrower and tubeshape. It also lacks the prominent crista dorsalis present in most variations of P. lundbeckii. Saether (1969) suggested that the synonymy of both species cannot be excluded. However, the two species can be separated based on partial COI DNA sequences (DNA barcodes)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on pages 20-22, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","Sublette, J. E. 1967. Type specimens of Chironomidae (Diptera) in the Cornell University collection. - Journal of the Kansas Entomological Society 477 - 564."]}

Rheocricotopus (Rheocricotopus) pauciseta Saether, 1969 Adults: Male described by Saether (1969), and Makarchenko and Makarchenko (2005). Female is unknown. Immatures: Pupa and larva described by Saether (1969) and in key by Epler (2001). Ecology and Habitat: Lotic habitats (Hudson et al. 1990). NE: Canada (Alberta, British Columbia, Labrador); USA (North Carolina). In the Palearctic Far East Russia, recently from China (Sichuan Province), Tibet by Liu et al. (2014b)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 19, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","Makarchenko, E. A. and Makarchenko, M. A. 2005. Chironomidae of the genus Rheocricotopus Thienemann et Harnisch, 1932 (Diptera, Chironomidae, Orthocladiinae) of the Russian Far East. - Eurasian Entomological Journal 4 (2): 126 - 134.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p.","Hudson, P. L. Lenat, D. R., Caldwell, B. A. and Smith D., 1990. Chironomidae of the southeastern United States: a checklist of species and notes on biology, distribution, and habitat. US Fish and Wildlife Publications 53 p.","Liu, W, Song, C. and Wang, X. 2014 b. Review of the subgenus Rheocricotopus (s. str.) Brundin, 1956 from China (Diptera, Chironomidae). - Pan-Pacific Entomologist 90: 100 - 106."]}

Rheocricotopus (Psilocricotopus) conflusirus Saether, 1985 Adults: Male described by Saether (1985). Female is unknown. Immatures: Unknown. Ecology and Habitat: Adults collected near reservoirs (Saether 1985). NE: USA (South Carolina)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 19, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948

Parametriocnemus graminicola (Lundbeck, 1898) Adults: Male described by Saether (1969) and Sublette (1966). Female is unknown. Immatures: Unknown. Ecology and Habitat: Adults collected near lentic habitats (Saether 1969). NE: Canada (Alberta, Northwest Territories, Yukon Territory); Greenland. In the Palearctic, recorded only in Far East Russia., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 17, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","Sublette, J. E. 1966. Type specimens of Chironomidae (Diptera) in the U. S. National Museum. - Journal of the Kansas Entomological Society 39 (4): 580 - 667."]}

Rheocricotopus (Psilocricotopus) glabricollis (Meigen, 1830) Adults: Male described by Lehman (1969), Saether (1985), Makarchenko and Makarchenko (2005), hypopygium in key by Pinder (1978). Female described by Saether (1985). Immatures: Pupa described by Saether (1985), figs. in Lehman (1969) under R. gouini, and in key by Langton (1991). Larva described in Namayandeh (2016), and in key by Epler (2001). Ecology and Habitat: Larvae inhabit lotic habitats (Hudson et al. 1990). NE: Canada (Manitoba, New Brunswick); USA (Georgia, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee). Widespread in the Palearctic., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 18, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Makarchenko, E. A. and Makarchenko, M. A. 2005. Chironomidae of the genus Rheocricotopus Thienemann et Harnisch, 1932 (Diptera, Chironomidae, Orthocladiinae) of the Russian Far East. - Eurasian Entomological Journal 4 (2): 126 - 134.","Pinder, L. C. V. 1978. A key to adult males of the British Chironomidae (Diptera), Vol. 1, The key; Vol. 2, Illustrations of the Hypopygia. - Freshwater Biological Association Scientific Publication 37: 1 - 169.","Langton, P. H. 1991. A key to pupal exuviae of West Palaearctic Chironomidae. Private publication. Printed by Graytones, UK. 386 p.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p.","Hudson, P. L. Lenat, D. R., Caldwell, B. A. and Smith D., 1990. Chironomidae of the southeastern United States: a checklist of species and notes on biology, distribution, and habitat. US Fish and Wildlife Publications 53 p."]}

Rheocricotopus (Rheocricotopus) amplicristatus Saether, 1985 Adults: Male and female described by Saether (1985). Immatures: Unknown. Ecology and Habitat: Adults collected near creeks (Saether 1985). NE: USA (Georgia, South Carolina)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 19, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948

Rheocricotopus (Rheocricotopus) tuberculatus Caldwell, 1984 Adults: Male and female described by Caldwell (1984) and Saether (1985). Immatures: Pupae described by Caldwell (1984) and Saether (1985). Larva by Caldwell (1984), Namayandeh et al. (2012), and in key by Epler (2001). Ecology and Habitat: In Georgia and North Carolina larvae occurred in second and third order piedmont streams feeding on detritus and diatoms (Caldwell, 1984). Namayandeh et al. (2012) collected the larvae from leaf litter in headwater streams located on the Precambrian Shield. NE: Canada (Ontario); USA (Florida, Georgia, North Carolina, South Carolina, Tennessee). Descriptions, Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on pages 19-20, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Caldwell, B. A. 1984. Two new species and records of other Chironomids from Georgia (Diptera: Chironomidae) with some observation on ecology. - Georgia Journal of Science 42: 81 - 96.","Namayandeh, A., Bilyj, B., Beresford, D. V., Somers, K. M. and Dillon P. J. 2012. Chironomidae (Diptera) larvae of Precambrian Shield headwater streams, Canada. - Zootaxa 3324: 1 - 165.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p."]}

Rheocricotopus (Psilocricotopus) chalybeatus (Edwards, 1929) Adults: Male described by Lehman (1969), Saether (1985), hypopygium in key by Pinder (1978). Female described by Saether (1985). Immatures: Pupa described by Saether (1985), in key by Langton (1991), figures in Lehman (1969). Larva by Cranston (1982). Ecology and Habitat: Larvae inhabit springs and streams (Cranston 1982, Lehman 1971). NE: Canada (Nunavut). Widespread in the Palearctic., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 18, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Pinder, L. C. V. 1978. A key to adult males of the British Chironomidae (Diptera), Vol. 1, The key; Vol. 2, Illustrations of the Hypopygia. - Freshwater Biological Association Scientific Publication 37: 1 - 169.","Langton, P. H. 1991. A key to pupal exuviae of West Palaearctic Chironomidae. Private publication. Printed by Graytones, UK. 386 p.","Cranston, P. S. 1982. A key to the larvae of the British Orthocladiinae (Chironomidae). - Scientific Publication Freshwater Biological Association 45: 1 - 152."]}

Key to the known Nearctic adult male Rheocricotopus Thienemann and Harnisch (Modified from Saether, 1985) Abbreviations: AnP = Anal Point; CD = Crista Dorsalis; Gc = Gonocoxite; Gs = Gonostyle; HP = Humeral Pit; IVo = Inferior Volsella; SVo = Superior Volsella. 1a. Gs either with prominent preapical triangular CD or it’s bent distally, and CD is more apical. SVo broadly rounded, never with projection. Subgenus Psilocricotopus..................................................... 2 1b. Gs either without apparent CD or its long, low and rounded distally. SVo with or without caudomedian projection. Subgenus Rheocricotopus...... 6 2a. HP of thorax small and indistinct. (Saether 1985, fig. 2B). AR R. (P.) conflusirus Saether 2b. HP of thorax large and distinct.AR> 0.7.......... 3 3a. HP very large, rectangular (Saether 1985, fig. 11B). Gs is not bent upwards distally (Saether 1985, fig. 11D)........... R. (P.) glabricollis (Meigen) 3b. HP moderately large, ovoid or circular, if very large and rectangular then Gs is bent upwards distally.................................................................... 4 4a. Gs with CD tooth-like located apically next to the megaseta (Lehmann 1969, fig. 1; Pinder 1978, fig. 38c).................. R. (P.) chalybeatus (Edwards) 4b. Gs with triangular preapical CD distinctly separatedfrom megaseta............................................... 5 5a. Gs strongly bent distally (Saether 1969, fig. 44) Costa not produced............................................................................. R. (P.) robacki (Beck and Beck) 5b. Gs not strongly bent (Saether 1985, fig. 4D). Costa moderately produced (Saether, 1985, fig. 4C).................................. R. (P.) chapmani (Edwards) 6a. SVo with or without caudomedian projection........................................................................ 7 6b. SVo broadly rounded without caudomedian projection.............................................................. 9 7a. SVo triangular without distinct caudomedian projection (Saether 1971, fig. 8D). IVo distally di- vided into two lobes. (Saether 1969, fig.47)...................................................... R. (R.) pauciseta Saether 7b. SVo with distinct caudomedian projection. IVo simple.................................................................... 8 8a. HP small (Fig. 2c). SVo with long finger-like caudomedian projections that meet medially (Fig. 3b–c)......................... R. (R.) reduncusoides sp. n. * 8b. HP large. SVo conical with short tapered caudomedian projections (Saether 1985, figs. 18b, d & e; Saether and Schnell 1988, figs. 3b & d).......... R. (R.) effusoides (Walker), R. (R.) effusus Saether, R. (R.) unidentatus Saether and Schnell † 9a. CD weak to absent. AR = 0.66–0.97. Costa with definite extension ≥ 15 μm.......................... 10 9b. CD present as long distally rounded ridge (Saether 1985, fig.16C). AR ≥ 1.0. Costa extension barely indicated R. (R.) amplicristatus Saether 10a. Gc with inner proximal margin bulges slightly before it meets the IVo that strongly projects medially (Saether 1969, fig. 43; Makarchenko and Makarchenko 2005, fig. 16). Costal extension ~ 30 μm............................. R. (R.) eminellobus Saether ‡ 10b. Gc with inner proximal margin continuing straight where it meets the IVo which ends in a small posteriorly directed lobe. (Saether 1985, fig.15C; Caldwell 1984, fig. 2). Costal extension 15–30 μm............. R. (R.) tuberculatus Caldwell ‡ *Will also key to R. reduncus Saether and Schnell, a Palearctic species. See diagnosis and remarks under R. reduncusoides, and Table 3 to separate the two species. †Theses three species in the effusus group are difficult to separate. See Table 3 for combination of characters, distinguishing the species in the effusus group. ‡ Adult male of R. eminellobus and R. tuberculatus are very similar. Key above can partially separate the two species. Female, pupa and larva of two species are quite distinguishable., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on pages 26-27, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Lehmann, J. 1969. Die europaischen Arten der Gattung Rheocricotopus Thien. und Harn. und drei neue Artvertreter dieser Gattung aus der Orientalis (Diptera, Chironomidae). - Archiv fur Hydrobiologie 66 (3): 348 - 381.","Pinder, L. C. V. 1978. A key to adult males of the British Chironomidae (Diptera), Vol. 1, The key; Vol. 2, Illustrations of the Hypopygia. - Freshwater Biological Association Scientific Publication 37: 1 - 169.","Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","Saether, O. A. 1971. Notes on general morphol- ogy and terminology of the Chironomidae (Diptera). - The Canadian Entomologist 103: 1237 - 1260. DOI: https: // doi. org / 10.4039 / Ent 1031237 - 9","Saether, O. A. and Schnell, O. A. 1988. Two new species of the Rheocricotopus (R.) effusus","Makarchenko, E. A. and Makarchenko, M. A. 2005. Chironomidae of the genus Rheocricotopus Thienemann et Harnisch, 1932 (Diptera, Chironomidae, Orthocladiinae) of the Russian Far East. - Eurasian Entomological Journal 4 (2): 126 - 134.","Caldwell, B. A. 1984. Two new species and records of other Chironomids from Georgia (Diptera: Chironomidae) with some observation on ecology. - Georgia Journal of Science 42: 81 - 96."]}

Rheocricotopus (Rheocricotopus) eminellobus Saether, 1969 Adults: Male described by Saether (1969), and Makarchenko and Makarchenko (2005). Female described by Saether (1969). Immatures: Pupa described by Saether (1969). Larva described by Saether (1985), in key by Epler (2001), and Namayandeh et al. (2012, 2016). Ecology and Habitat: Larvae are shredders and inhabit leaf litter in running waters (Namayandeh et al. 2012). NE: Canada (Alberta, Labrador, Ontario); USA (North Carolina, Ohio, South Carolina, Tennessee). In the Palearctic, found only in the Far East Russia., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 19, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","Makarchenko, E. A. and Makarchenko, M. A. 2005. Chironomidae of the genus Rheocricotopus Thienemann et Harnisch, 1932 (Diptera, Chironomidae, Orthocladiinae) of the Russian Far East. - Eurasian Entomological Journal 4 (2): 126 - 134.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p.","Namayandeh, A., Bilyj, B., Beresford, D. V., Somers, K. M. and Dillon P. J. 2012. Chironomidae (Diptera) larvae of Precambrian Shield headwater streams, Canada. - Zootaxa 3324: 1 - 165."]}

Parametriocnemus hamatus (Johannsen, 1934) (Figs 1a–e) Material examined. Parametriocnemus (1 ♂), Old Man Joe Mt. (No such location was found in Québec), Québec, 26 April 1968, Collected by D. R. Oliver, No. CH 767 Diagnostic characters. Virga with 2–3 long narrow branches. AnP with inflated basal 1/2 to 2/3 with 4 stout setae on each side. IVo narrow and apically hooked. Gc with large projecting CD. Male (n=1). Total length = 3.2 mm Coloration of slide-mounted specimen: Male head and thorax light brown, abdomen golden-brown, halter hyaline to slightly golden, and wing golden-brown. Head. Antenna (Fig. 1a), with 13 flagellomeres, ultimate flagellomere (L = 606 μm) with apical setae (L = 35 μm); AR = 1.3. Eyes bare, with parallelsided dorsomedial extension (Fig. 1b). Tentorium L = 147 µm (Fig. 1b). Four coronal setae, six orbitals in single row on each side, Clypeus squared (L = 64 µm, W = 68 µm), bearing nine setae. Palpomeres p 1–5, lengths (µm): 36, 42, 141, 143, 207; p 3 with 2 sensilla chaetica. Thorax. As in Fig. 1c. Antropronotals 11, uniserial; prealars 6, uniserial. Scutum with six setae in single row. Wing. Wing with fine punctation and hairs covering most of membrane (Fig. 1d). L = 2.1 mm, W = 0.6 mm. R with 24 setae, R 1 with 15, R 4+5 with 60 setae. Costa extends pass R 4+5. R 4+5 distal to M 3+4. Cu curved. Legs. Fore legs with two spurs (Ls = 42 µm, 30 µm), mid tibia with two spurs (Ls= 34 µm, 24 µm) and hind tibia with 2 spurs (Ls = 59 µm, 19 µm) and comb with 12–13 stout setae. Pseudospurs are absent. Mid and hind femurs with keel. Pulvilli reduced. Lengths and proportions of legs in Table 1. Hypopygium. As in Fig. 1e. Segment IX bilobed. Virga present with 2–3 long narrow branches. Anal point L = 61 µm. Superior volsella tapered. Inferior volsella narrow and apically hooked-shaped. Gonocoxite longer than wide (L = 203 µm, W = 85 µm) with cluster of long setae medially just below the inferior volsella. Gonostyle more or less rectangular; crista dorsalis large triangular-shaped and strongly projecting above dorsal margin; megaseta L = 13 µm. HR = 3.8, HV = 2.4. Remarks. The species was first described by Johannsen, 1934 as Metriocnemus hamatus and indicated its close relationship with Meteriocnemus lundbecki Johannsen differing from the latter only in coloration and hypopygium. Sublette (1967) re-described the species as Paraphaenocladius hamatus (Joh.) with illustration of the male hypopygium., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 20, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Johannsen, O. A. 1934. New species of North American Ceratopogonidae and Chironomidae. - Journal of the New York Entomological Society 42 (3): 343 - 352.","Sublette, J. E. 1967. Type specimens of Chironomidae (Diptera) in the Cornell University collection. - Journal of the Kansas Entomological Society 477 - 564."]}

Namayandeh, Armin, Beresford, David V. (2018): A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract. CHIRONOMUS Journal of Chironomidae Research 31: 16-29, DOI: 10.5324/cjcr.v0i31.2531

Rheocricotopus (Rheocricotopus) reduncusoides sp. n. h t t p: / / z o o b a n k. o rg / u r n: l s i d: z o o b a n k. o r g: a c t: B D 1 A E E 5 9 - F 9 A 2 - 4 E B B - 9 B B 6 - 791096FFB959 (Figs 2a–c & 3a–c) Material examined. Holotype: 1 ♂ slide-mounted in Canada Balsam, Bathurst Island, Nunavut, 75° 43’ N, 98° 25’ W, 10 July 1969, Coll. H.V. Danks, CH 1214 (CNC). Paratypes: 4 ♂♂ slide-mounted in Canada Balsam, Bathurst Island, Nunavut, 75° 43’ N, 98° 25’ W, 10 July 1969, Coll. H.V. Danks, CH 1214 (CNC). Rheocricotopus (Rheocricotopus) reduncus Saether and Schnell, 1988: 3 Paratypes, ♂♂, Jostedøla River at Inlet to small lake, Luster, Sogn and Fjordane, Norway, 23/7’ 86, A. Fjellheim and A. Schnell (ZMB NO. 116). Etymology: From Latin, reduncus meaning curved or hooked backward which refers to the shape of caudomedian projection of superior volsella, and the epithet of the most closely related species, and New Latin oides, referring to the likeness of form. Diagnostic characters. HP small to indistinct. SVo with long caudomedian projections evenly curved, medially projected and opposing each other, apex slightly pointed. Sternapodme broad horizontal band. IVo simple, small, blunt, slightly dilated at the apex. AR = 0.75. BR 1–3 = 1.4, 1.2, 1.4. AnP 67 µm long with 13–19 setae. Gs with slight bent distally, CD developed, long and low. HR = 1.6. Male (n = 5). Total length = 3.1mm Coloration of slide-mounted specimen: Head and thorax dark brown, abdomen golden-brown, halter light brown with dorsolaterals darker, and wing yellowish brown. Head. As in Fig. 2a. Antenna (Fig. 2b) with 13 flagellomeres, ultimate flagellomere 332 μm long, AR = 0.72–0.79 (0.75). Tentorium L = 152 µm. 4 Coronal setae. Temporal setae consisting of: 3 postorbitals, 1 inner verticals, 2 outer verticals present. Clypeus wider than long (L = 119 µm, W = 132 µm), bearing six setae. Palpomere p 1–5 lengths (µm): 51, 55, 79, 88, 144; sensilla clavata difficult to see on 3 rd palpomere. Thorax. As in Fig. 2c. 7–8 antepronotals. 4–5 (4) acrostichals close to antepronotum (L = 11 µm). Dorsocentrals 7–9, uniserial (Ls = 57–69 µm). Prealars 2–3. Wing. As in Fig. 3a. Wing with fine punctation. L = 2 mm, W = 0.5 mm. Squama with 4–5 setae, brachiulum with 1 seta, R with 5–6 setae, R 1 bare, R 4+5 with 2 setae. Costa slightly extends past R 4+5 (L = 30 µm). Legs. Fore legs with tibial spur 41 µm long, mid tibial spurs 25 and 16 µ long, hind tibial spurs 44 and 23 µ long, comb with 15–17 setae. Pseudospurs are absent on t 1 and t 2 of mid and hind legs. Mid and hind femur with keel. Pulvilli well-developed, almost half as long as the claws. Lengths and proportions of legs in Table 2. Hypopygium. As in Fig. 3b. Laterosternite IX with 6–7 setae (L = 25 µm). Phallapodme (59–66) 64 μm long; sternapodeme (90–105) 99 μm long. Anal point mostly hyaline, triangular with 13–19 setae, (56–89) 67 µm long. Superior volsella (Fig. 3b–c) with caudomedian projections long, finger-like, curved evenly before meeting medially, L = 84 µm. Inferior volsella simple triangular, lobe with blunt tip and slightly dilated at apex, L = 19 µm. Gonocoxite 278 µm long. Gonostyle curved with slight bent distally (Fig. 3b), L = 171 µm. Crista dorsalis long and low, megaseta L = 17 µm. HR = 1.6, HV = 1.8. Remarks. This species is closely related to the Rheocricotopus (Rheocricotopus) reduncus. Combination of lower AR, lower BR, higher number of setae on AnP, longer AnP, a more evenly curved caudomedian projection of SVo (Figs 3b–c; compared to mainly bent in R. reduncus Fig. 4a–b), apically dilated IVo, broad horizontal sternapodeme (Fig. 3b; compared to narrow arched in R. reduncus, Fig. 4c), lower HR and HV separates the two species. Based on the form of SVo this species belongs to the effusus group. Given the similarity of R. reduncus, R. reduncuiodes with other species within the effusus group a re-evaluation of the distinguishing characters is required to separate the species. In the effusus group the anal point length and thoracic chaetotaxy characters may not be sufficient to separate the species in this group. Saether and Schnell (1988) provided a somewhat better alternative for separating species in this group by listing the main comparative characters in a table. Following their example, we modified and corrected some of these characters and added few more (Table 3). A notable change to the characters given by Saether and Schnell (1988) is the AR range of R. reduncus. The Norwegian specimens of R. reduncus described by Saether and Schnell (1988) have AR 0.83–0.90 while Makarchenko and Makarchenko (2005) described the species from Far Eastern Russia with AR range of 1.16–1.17. This somewhat changes the state of characters given by Saether (1985) and discussed by Saether and Schnell (1988). If we are to consider Makarchenko and Makarchenko (2005) description of R. reduncus then the trend 8 of Saether’s (1985) classification of the genus, describing a male AR of 0.6–0.8, does not longer hold for this species. Consequently, the characteristic of AR should no longer be considered apomorphic for R. reduncus. For 1 st of trend 7 in Saether’s (1985) both R. reducncus and R. reduncusoides are synapomorphous (digitiform caudomedian projection) and symplesiomorphous for second (with small humeral pit). The broad horizontal sternapodeme of R. reduncusoides is autapomorphic for this species., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on pages 22-26, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. and Schnell, O. A. 1988. Two new species of the Rheocricotopus (R.) effusus","Makarchenko, E. A. and Makarchenko, M. A. 2005. Chironomidae of the genus Rheocricotopus Thienemann et Harnisch, 1932 (Diptera, Chironomidae, Orthocladiinae) of the Russian Far East. - Eurasian Entomological Journal 4 (2): 126 - 134."]}

Rheocricotopus (Psilocricotopus) robacki (Beck and Beck, 1964) Adults: Male described by (Beck and Beck 1964) as Trichocladius robacki and by Saether (1969) as R. kenorensis. Female described by Saether (1985). Immatures: Pupa and larva described by Saether (1985). Larva in key by Epler (2001). Ecology and Habitat: Fast flowing streams (Saether 1969). NE: Canada (Alberta, British Columbia, Ontario, Saskatchewan, Yukon Territory); USA (Alabama, Arizona, California, Florida, Georgia, Mississippi, Montana, New York, North Carolina, Ohio, Pennsylvania, South Carolina, South Dakota, Tennessee). Recently reported from the Palaearctic China (Xinjiang Uyghur Auto. Region) by Liu et al. (2014a)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 18, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Beck, W. M. and Beck, E. C. 1964. New Chironomidae from Florida (Diptera). - The Florida Entomologist 47: 201 - 207.","Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154.","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. Special Publication SJ 2001 - SP 13. North Carolina Department of Environment and Natural Resources Division of Water Quality and St. Johns River Water Management District, Palatka, FL, 530 p.","Liu, W., Lin, X. and Wang, X. 2014 a. A review of Rheocricotopus (Psilocricotopus) chalybeatus species group from China, with description of three new species (Diptera, Chironomidae). - ZooKeys 388: 17 - 34."]}

Parametriocnemus eoclivus Saether, 1969 Adults: Male and female described by Saether (1969). Immatures: Pupa and larva described by Saether (1969). Ecology and Habitat: Larvae inhabit lotic habitats (Saether 1969). NE: Canada (Québec); USA (North Carolina, Tennessee). In Palearctic, only recorded in Italy (dubious record)., Published as part of Namayandeh, Armin & Beresford, David V., 2018, A NEW SPECIES IN THE RHEOCRICOTOPUS (R.) EFFUSUS GROUP FROM CANADA WITH A REVIEW OF THE NEARCTIC SPECIES OF RHEOCRICOTOPUS AND PARAMETRIOCNEMUS (CHIRONOMIDAE: ORTHOCLADIINAE) Abstract, pp. 16-29 in CHIRONOMUS Journal of Chironomidae Research 31 on page 17, DOI: 10.5324/cjcr.v0i31.2531, http://zenodo.org/record/7994948, {"references":["Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). - Bulletin of Fisheries Research Board of Canada 170: 1 - 154."]}