Research Article |
Jade Savage‡, Vera S. Sorokina§
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Corresponding author: Jade Savage ( jsavage@ubishops.ca ) Academic editor: Pierfilippo Cerretti
© 2021 Jade Savage, Vera S. Sorokina.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Savage J, Sorokina VS (2021) Review of the North American fauna of Drymeia Meigen (Diptera, Muscidae) and evaluation of DNA barcodes for species-level identification in the genus. ZooKeys 1024: 31-89. https://doi.org/10.3897/zookeys.1024.60393
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Abstract
The North American fauna of Drymeia Meigen was studied. Four new species are described (Drymeia hucketti sp. nov., Drymeia ponti sp. nov., Drymeia vockerothi sp. nov., Drymeia woodorum sp. nov.), and three new synonymies are proposed: Drymeia amnicola (Huckett, 1966) (= Drymeia rivalis (Huckett, 1966), syn. nov.); Drymeia glacialis (Rondani, 1866) (= Drymeia alpicola (Rondani, 1871), syn. nov.); and Drymeia spinitarsis (Aldrich, 1918) (= Drymeia longiseta Sorokina & Pont, 2015, syn. nov.). An annotated checklist, DNA barcodes (when available), and keys for each sex of the 24 named species of North American Drymeia are provided. The utility of DNA barcodes for the identification of Drymeia species across a wide geographical range was explored using sequences from five countries. A match between morphology and DNA barcodes was found for 71% (22/31) of species studied (including three unnamed taxa). The remaining nine species clustered into two groups of taxa with very little interspecific variation within clusters (groups of two and seven species). Caution is advised against using DNA barcoding as the only determination tool for Drymeia material without prior knowledge of its limitations for certain species groups.
Keywords
Azeliinae, Barcoding Index Number (BIN), checklist, identification key, North America
Introduction
The genus Drymeia Meigen, 1826 includes more than 130 species of mid-sized muscids that can usually be distinguished from other members of the Muscidae by the combination of a dilated and densely setose gena, proclinate orbital bristles in the female, and three or more posterodorsal bristles on the hind tibia. An apical ventral spur of variable size can be found on the hind tibia of males in many species (see Fig.
Drymeia is entirely restricted to the northern hemisphere and most species have an alpine and/or northern distribution. Drymeia segnis (Holmgren, 1883), for example, is among the few insect species found as far north as Qaanaaq (formerly known as Thule), Greenland (
While these flies can be collected using various passive collecting devices, net sweeping will often yield a good range of species since many can be found basking on sunny rocks or visiting flowers. Many species of Drymeia are known to consume pollen in addition to nectar, and
Not much is known of the immature stages of Drymeia. Some larvae can be found in humus soil or cow dung and
The vast majority of North American species were originally described under Pogonomyia Rondani and, to a lesser extent, Bebryx Gistl, Trichopticoides Ringdahl, Eupogonomyia Malloch, and Pogonomyioides Malloch before these taxa were synonymised with Drymeia by
Until recently, only Drymeia alpicola (Rondani, 1871) and D. segnis were recorded as having holarctic distributions. The works of
All taxonomic contributions to the North American fauna of Drymeia to date have focused on local fauna, making it difficult to identify specimens outside of these areas. Consequently, the first objective of this work was to develop comprehensive and well-illustrated keys to both sexes of all North American species of Drymeia. We also aimed to produce an annotated checklist of these species (including new synonymies and updated distribution ranges) and describe four new species. Finally, we explored the utility of DNA barcodes and Barcoding Index Number (BIN) assignments (
Materials and methods
Drymeia specimens examined in this study are housed in the Academy of Natural Sciences of Philadelphia, Philadelphia, USA (
Morphology
Morphological terms follow
The following abbreviations are used in the text:
a anterior;
acr acrostichal;
ad anterodorsal;
av anteroventral;
dc dorsocentral;
p posterior;
pd posterodorsal;
pv posteroventral;
v ventral;
F femur;
T tibia.
Label data of primary type material examined are presented verbatim, with “/” indicating a change of line and “;” a change in label.
DNA barcoding
The Barcode Index Number (BIN) system uses the Refined Single Linkage (RESL) algorithm to group COI sequences in the Barcode of Life Data System (BOLD) (http://boldsystems.org) into genetic clusters (BINs) representing proxies for species (
All DNA barcodes (Folmer region of COI gene) were sequenced at the University of Guelph Biodiversity Institute of Ontario following protocols published in
In the public dataset "Drymeia specimens examined for Savage and Sorokina 2021" (dx.doi.org/10.5883/DS-DRYNEW) available on BOLD. Specimen details, including BIN assignment, sex, and GenBank accession numbers for this data set can also be found in Suppl. material
Results
Drymeia
Drymeia Meigen, 1826: 204. Type species: Drymeia obscura Meigen, 1826 [= Musca hamata Fallén, 1823] (monotypy).
Drymia Agassiz, 1847: 130. Unjustified emendation of Drymeia Meigen.
Eriphia Meigen, 1826: 206. [Junior homonym of Eriphia Latreille, 1817: Crustacea] Type species: Eriphia cinerea Meigen, 1826 (monotypy).
Bebryx Gistl, 1848: ix. [Replacement name for Eriphia Meigen, 1826] Type species: Eriphia cinerea Meigen, 1826 (automatic).
Pogonomyia Rondani, 1871 [1870]: 336. Type species: Pogonomyia alpicola Rondani, 1871 (monotypy).
Neoeriphia Schnabl & Dziedzicki, 1911: 195 [as subgenus of Eriphia Meigen, 1826]. Type species: Eriphia metatarsata Stein, 1907 (monotypy).
Neopogonomyia Schnabl & Dziedzicki, 1911: 198 [as subgenus of Pogonomyia Rondani, 1871]. Type species: Aspilia brumalis Rondani, 1866, by designation of Séguy, 1923, Faune Fr., 6: 295. [A. brumalis is listed by Schnabl and Dziedzicki as a junior synonym of meadei, and meadei is the first of two included species.]
Pogonomyioides Malloch, 1919: 67. Type species: Pogonomyioides atrata Malloch, 1919 [= Aricia segnis Holmgren, 1883] (original designation).
Eupogonomyia Malloch, 1921: 178. Type species: Eupogonomyia pribilofensis Malloch, 1921 (original designation).
Trichopticoides Ringdahl, 1931: 173. Type species: Musca decolor Fallén, 1824 [= Musca vicanus Harris, 1780] (monotypy).
Remarks
While the exact systematic position of Drymeia remains debated (
Key to males of North American species of Drymeia Meigen
(males of D. latifrons (Malloch) unknown)
| 1 | Calypter black. Recorded only from Mexico | 2 |
| – | Calypter pale brown to yellow. Not recorded from Mexico | 3 |
| 2 | Frons at narrowest point at least 1.5 × as wide as ocellar triangle and with frontal vitta widely exposed (Fig. |
ponti sp. nov. |
| – | Frons narrower than ocellar triangle and with fronto-orbital plates usually touching at narrowest point (Fig. |
aterrima (Wulp) |
| 3 | Anepimeron haired | 4 |
| – | Anepimeron bare | 5 |
| 4 | Epandrium clotted with very long curly hair forming a distinctive brush at tip of abdomen (Fig. |
chillcotti (Huckett) |
| – | Epandrium with regular setae; fore coxa prolonged into a knob-like structure (Fig. |
segnis (Holmgren) |
| 5 | Prealar reduced or absent; if visible, then no longer than 1/2 the length of posterior notopleural bristle; 2+4 dc | 6 |
| – | Prealar strong, approximately as long as posterior notopleural bristle; dc variable | 12 |
| 6 | Lower margin of face strongly projecting beyond lower level of frons (Fig. |
pribilofensis (Malloch) |
| – | Lower margin of face not or only weakly projecting beyond lower level of frons; F2 not as above; outer claw of fore tarsus not strongly enlarged (sometimes slightly larger in D. setibasis) | 7 |
| 7 | T3 with ventral apical process very strongly developed, longer than width of hind tibia (Fig. |
groenlandica (Lundbeck) |
| – | T3 with apical process short to well-developed but always much shorter than maximum width of hind tibia (Fig. |
8 |
| 8 | F2 with 1 or 2 short stout ventral spines near middle (Fig. |
cantabrigensis (Huckett) |
| – | F2 without short stout spines near middle but sometimes with short ventral spines closer to base | 9 |
| 9 | F2 with a very strong clump of curved av near the very base (Fig. |
setibasis (Huckett) |
| – | F2 at most with a few long delicate av on basal 1/3 (Fig. |
10 |
| 10 | Fore coxa with a patch of 4 or 5 very long, strong apical bristles, at least as long as length of coxa; smaller species (5.5 mm) | woodorum sp. nov. |
| – | Fore coxa without long bristles, only with short spines; larger species (7.7–9.5 mm) | 11 |
| 11 | Frons broad with black frontal vitta exposed and at least 2 × as wide as fore ocellus (Fig. |
neoborealis (Snyder) |
| – | Frons narrow with fronto-orbital plates touching at narrowest part (Fig. |
vockerothi sp. nov. |
| 12 | Parafacial broad in lateral view, wider than length of first flagellomere midpoint (Fig. |
13 |
| – | Parafacial in lateral view no wider than width of first flagellomere midpoint (Fig. |
15 |
| 13 | Frons broad with black frontal vitta exposed and at least as wide as fore ocellus at narrowest point; fore and mid coxa with very long fine hair, crinkled near apex; mid tarsomere 1 with spines in av and pv rows long, some longer than width of tarsomere (Fig. |
spinitarsis (Aldrich) |
| – | Frons narrow with fronto-orbital plates touching at narrowest point; hairs of fore and mid coxa straight, not as above; mid tarsomere 1 with spines in av and pv rows shorter than width of tarsomere; T3 with ventral apical process present or absent | 14 |
| 14 | F2 with complete row of av and with 2 or 3 long and strong v about 1/3 from apex (Fig. |
quadrisetosa (Malloch) |
| – | F2 with complete row of av but without long and strong v about 1/3 from apex (Fig. |
glacialis (Rondani) |
| 15 | Wing with membrane and veins conspicuously yellow over most of the surface; T3 with a complete row of numerous short, sub-equal ad, no longer than diameter of tibia | flavinervis (Malloch) |
| – | Wing usually hyaline or infuscated brown but if yellowish, then row of ad on T3 not as above | 16 |
| 16 | F2 with av and pv rows short and irregular, restricted to apical 1/2 and with bristles no longer than diameter of femur (Fig. |
similis (Malloch) |
| – | F2 with av and/or pv rows variable, but always with some (or many) bristles longer than basal diameter of femur (as in Fig. |
17 |
| 17 | F2 with both av (Fig. |
18 |
| – | F2 with either av row incomplete (as in Fig. |
19 |
| 18 | Prementum dusted; scutum dusted, abdomen grey dusted except for most of tergite I+II, a wide central patch on tergite III and central vittae on tergites IV and V (Fig. |
firthiana (Huckett) |
| – | Prementum undusted, shiny; scutum and abdomen mostly shiny; F2 and/or F3 usually with a few long fine and slightly crinkled p-pv on basal 1/4 (Fig. |
profrontalis (Huckett) |
| 19 | F2 with pv row incomplete, with 4–7 very long pv on apical 1/2, these at least 2 × as long as width of femur | 20 |
| – | F2 with pv row complete, with some bristles longer than width of femur at least on basal 1/2 (Fig. |
21 |
| 20 | Lower margin of face projecting slightly beyond lower level of frons (Fig. |
hucketti sp. nov. |
| – | Lower margin of face not projecting beyond lower level of frons (Fig. |
minor (Malloch) |
| 21 | F2 with pv row sparse and delicate, with only a few bristles longer than width of femur (Fig. |
santamonicae (Huckett) |
| – | F2 with pv row regular and thicker than above, with many bristles longer than width of femur on basal 2/3 (Fig. |
22 |
| 22 | T2 without ad on apical 1/3, California records doubtful (see remarks under D. aldrichi) | aldrichi (Malloch) |
| – | T2 usually with 1 or 2 ad present on apical 1/3 (these sometimes absent), recorded only from California | amnicola (Huckett) |
Key to females of North American species of Drymeia Meigen
| 1 | Prealar bristle present and strong, as long as 2nd notopleural bristle | 2 |
| – | Prealar bristle absent or short, always much shorter than 2nd notopleural bristle | 19 |
| 2 | Parafacial in lateral view broader than width of first flagellomere (Fig. |
3 |
| – | Parafacial in lateral view variable; F2 not as above; 2+3 dc. Size variable but usually < 7.5 mm | 5 |
| 3 | Mid tarsomere 1 with spines in av and pv rows long, some longer than width of tarsomere (Fig. |
spinitarsis (Aldrich) |
| – | Mid tarsomere 1 with spines in av and pv rows shorter than 1/2 the width of tarsomere; head in lateral view no higher than wide (as in Fig. |
4 |
| 4 | Veins of basal region of wing except costa yellowish to pale brown | quadrisetosa (Malloch) |
| – | Veins of basal region of wing dark brown to yellowish | glacialis (Rondani) [Nearctic females of D. glacialis generally exhibit a darker wing base than those of D. quadrisetosa but females of these two species are difficult to distinguish (see Remarks under D. glacialis for additional details)] |
| 5 | Entire wing membrane and all veins rich yellow (costa brownish only near wing tip) (Fig. |
flavinervis (Malloch) |
| – | Wing membrane never rich yellow throughout most of the surface and most veins brownish near apex; parafacial shining or dusted | 6 |
| 6 | Prementum dusted and no longer than palpus; body light grey dusted; T2 with 2 or 3 ad; T3 with apical pv short or absent; parafacial and profrons in anterior view entirely dusted. | firthiana (Huckett) |
| – | Prementum shiny, usually longer than palpus; body mostly undusted and shiny except in D. santamonicae; T2 with ad variable; T3 with apical pv present or absent; dusting of parafacial and profrons variable | 7 |
| 7 | T2 with at least 1 av, costal spinules usually long and strong with costal spine at least 1.5 × as long as surrounding spinules (Fig. |
8 |
| – | T2 usually without av; costal spinules and costal spine variable | 10 |
| 8 | T3 with apical pv usually absent but if visible, then no longer than 1/2 the length of apical av; arista with longest hair no longer than base or arista; fore tarsomere 5 distinctly flattened (Fig. |
ponti sp. nov. (part, usually without av on T2) |
| – | T3 with apical pv present, at least 1/2 as long as apical av; arista with longest hair usually slightly longer than base of arista; fore tarsomere 5 at most slightly flattened (Fig. |
9 |
| 9 | Parafacial entirely dusted or with small undusted line near base of antenna (Fig. |
similis Malloch (part, usually without av on T2) |
| – | Parafacial with large undusted shiny patch near base of antenna (Fig. |
latifrons (Malloch) |
| 10 | Parafacial entirely dusted or with small undusted line near base of antenna (Fig. |
similis (Malloch) (part) |
| – | With different combination of characters | 11 |
| 11 | Fronto-orbital plates and ocellar triangle mostly undusted and shiny (Fig. |
profrontalis (Huckett) |
| – | Fronto-orbital plates mostly to completely dusted (Fig. |
12 |
| 12 | Parafacial in anterior view with large undusted shiny patch near base of antenna reaching up to or almost up to eye (Fig. |
13 |
| – | Parafacial in anterior view completely or mostly dusted, at most with a small shiny patch near base of antenna (Fig. |
15 |
| 13 | Scutum and abdomen dark grey with thin brownish grey pollen; ocellar triangle entirely dusted; parafacial in lateral view no broader than width of first flagellomere at midpoint; recorded only from California | santamonicae (Huckett) |
| – | Scutum and abdomen black and mostly shiny (Fig. |
14 |
| 14 | Parafacial in lateral view no broader than width of first flagellomere (as in Fig. |
amnicola (Huckett) (part) |
| – | Parafacial in lateral view broader than width of first flagellomere (Fig. |
hucketti sp. nov. |
| 15 | Fore tarsomere 5 moderately to distinctively flattened (Fig. |
16 |
| – | Fore tarsomere 5 not or only slightly flattened (Fig. |
17 |
| 16 | Costal spinules long and strong with costal spine 2 × as long as surrounding spinules (Fig. |
ponti sp. nov. (part) |
| – | Costal spinules not strongly developed, with costal spine only slightly larger than surrounding spinules (Fig. |
aldrichi (Malloch) |
| 17 | Arista with longest hair 1.5–2.0 × width of base of arista (Fig. |
aterrima (Wulp) |
| – | Arista with longest hair as long as base of arista (Fig. |
18 |
| 18 | Larger species, 6.2–7.3 mm; F2 usually with av reduced on apical 1/2, these no longer than width of femur; recorded only from California | amnicola (Huckett) (part) |
| – | Smaller species, 4.5–5.7 mm; F2 usually with 2 strong av on apical 1/2, longer than width of femur; broader distribution | minor (Malloch) |
| 19 | Anepimeron and/or katepimeron with hairs | 20 |
| – | Anepimeron and katepimeron both bare | 21 |
| 20 | F2 with 1 or 2 strong av setae on apical 1/2, as strong as or stronger than av near base; lower margin of face not projecting beyond lower level of frons; usually a larger species 7.4–9.3 mm. | chillcotti (Huckett) |
| – | F2 without or with only a short weak av in apical 1/2; lower margin of face slightly projecting beyond lower level of frons; usually a smaller species 6.0–8.5 mm | segnis (Holmgren) |
| 21 | Parafacial in lateral view narrow at midpoint, < 1/2 the width of first flagellomere (Figs |
22 |
| – | Parafacial in lateral view wide at midpoint, at least as wide as first flagellomere (Fig. |
23 |
| 22 | Large species (7.5–9.0 mm) (Fig. |
vockerothi sp. nov. |
| – | Small species (4.9–5.5 mm) (Fig. |
woodorum sp. nov. |
| 23 | T2 with av present | setibasis (Huckett) (part) |
| – | T2 with av absent | 24 |
| 24 | Lower margin of face not projecting beyond lower level of frons; prealar stronger than ground setulae but always much weaker and shorter than posterior notopleural bristle | 25 |
| – | Lower margin of face projecting beyond lower level of frons; prealar usually absent (a weak prealar sometimes visible in D. cantabrigensis) | 26 |
| 25 | F2 with longest av on apical 1/3 as long as width of femur; abdomen lightly dusted, subshiny | groenlandica (Lundbeck) |
| – | F2 with longest av on apical 1/3 shorter than width of femur; abdomen heavily dusted | neoborealis (Snyder) |
| 26 | Lower margin of face usually strongly projecting beyond lower level of frons (Fig. |
pribilofensis (Malloch) |
| – | Lower margin of face projecting only slightly beyond lower level of frons (Fig. |
setibasis (Huckett) (part), cantabrigensis (Huckett) [Some females of D. pribilofensis occasionally do not exhibit the full combination of characters listed in the key and may therefore be difficult to separate from D. setibasis and D. cantabrigensis.] |
Annotated checklist and new species descriptions
Drymeia aldrichi
Pogonomyia aldrichi Malloch, 1918: 281.
Pogonomyia unicolor, Stein, 1920: 22.
Type material examined
Pogonomyia aldrichi – Holotype male labelled “
Other material examined
2 females: Nearctic: USA: Idaho: Moscow (
Distribution
Nearctic: Canada (Alberta), USA (Washington to California (see remarks), Wyoming).
DNA barcode
None available.
Remarks
While California was listed in the distribution of D. aldrichi by
Drymeia amnicola
Pogonomyia amnicola Huckett, 1966: 291.
Pogonomyia rivalis Huckett, 1966: 293. syn. nov.
Type material examined
Pogonomyia amnicola – Holotype male labelled “Sardine Crk./Mono Co./ Elev. 8500/ Cal. VI-28-51”; “J.W. MacSwain/ Collector”; “Pogonomyia/ amnicola/ n. sp./ holotype [red]”; “California Academy/ of Sciences/ Type No. 10149” (
Paratypes: all with “Pogonomyia/ amnicola/ n. sp./ paratype [blue]”. 2 males, “Sardine Crk./Mono Co./ Elev. 8500/ Cal. VI-28-51”; “J.W. MacSwain/ Collector” (
Pogonomyia rivalis – Allotype female labelled “Sardine Crk./Mono Co./ Elev. 8500/ Cal. VI-28-51”; “S.M. Kappos/ Collector”; “Pogonomyia rivalis/ n. sp./ Allotype [orange]” (
Other material examined
1 male, 7 females: Nearctic: USA: California: Leavitt Mdw, 1 mi. S saddlebag Lake, Sardine Creek (
Distribution
Nearctic: USA (California).
DNA Barcode
None available.
Remarks
In his original descriptions,
Furthermore,
Drymeia aterrima
Pogonomyia aterrima Wulp, 1896: 335.
Type material examined
None.
Other material examined
More than 100 males and females: Neotropical: Mexico: Durango: 10 mi W. El Salto. (
Distribution
Nearctic: USA (Washington, Montana, but see remark below). Neotropical: Mexico (Durango).
DNA Barcode
None available.
Remarks
This species was recently redescribed by
Drymeia cantabrigensis
Eupogonomyia cantabrigensis Huckett, 1965a: 300.
Type material examined
Eupogonomyia cantabrigensis – Holotype male labelled “Cambridge Bay /N.W.T. 18.VII. 1950/ G. K. Sweatman”; “Type [red]”; “HOLOTYPE/ CNCNo. 8367 [red]”; “Eupogonomyia/ cantabrigensis Huck./ det. H.C. Huckett” (
Other material examined
More than 100 males and females: Nearctic: Canada: North West Territories: Arviat [formerly Eskimo point], Hooper Is., Mould Bay (Prince Patrick Is.), Sachs Harbour, Victoria Is.; Nunavut: Banks Is., Cambridge Bay, Kugluktuk, Masik Riv. (Banks I.); Yukon Territory: Herschel Is. (BUIC,
Distribution
Nearctic: Canada (Northwest Territories, Nunavut, Yukon Territory).
DNA Barcode
BOLD BIN: BOLD:AAD7664 (BIN merge with several other species, see Fig.
Remarks
While always lacking an anteroventral bristle on the midtibia, the females of this species are sometime indistinguishable from those of D. setibasis where this bristle is either present or absent. DNA barcodes for D. cantabrigensis were very similar and in some case identical to those of other species forming a cluster (or BIN merge) of seven named species (including D. setibasis) in BOLD:AAD7664 (Fig.
Drymeia chillcotti
Bebryx chillcotti Huckett, 1965a: 302.
Type material examined
Bebryx chillcotti – Holotype male labelled “Kidluit Bay, N./ Richards Is. W./ 27-VIII 1948. T./ J. R. Vockeroth”; “Type [red]”; “HOLOTYPE/ CNCNo. 8369 [red]”; “Bebryx/ chillcotti Huck./ det. H.C. Huckett” (
Other material examined
More than 180 males and females: Nearctic: Canada: British Columbia: Summit of Pink Mtn; Northwest Territories: 20 and 21 m. e. Tuktoyaktuk; Nunavut: Arviat [formerly Eskimo point], Char river nr Ranking Inlet, Clyde (Baffin Island), Coral Harbor (Southampton Island), Ford Lake, Landing Lake (7.5 km NW of Rankin inlet), Masik River (Banks Island.), Meliadine river, Padlei, Victoria Island; Yukon Territory: British Mts, Firth River, Richardson Mts, USA: Alaska: Noluk. Palaearctic: Russia: Taymyr Peninsula (BUIC,
Distribution
Nearctic: Canada (British Columbia, Northwest Territories, Nunavut, Yukon Territory), USA (Alaska). Palaearctic: Russia (Taymyr Peninsula).
DNA Barcode
BOLD BIN: BOLD:ACA8934. See Suppl. material
Remark
DNA barcodes for material from Canada and Russia (Fig.
Drymeia firthiana
Pogonomyia firthiana Huckett, 1965a: 298.
Type material examined
Pogonomyia firthiana – Holotype male labelled “Firth River, Y.T./ 14-VII-1956/E.F. Cashman”; “Type [red]”; “Type/ HOLO/ No. 8366 [red]”; “Pogonomyia/ firthiana Huck./ det. H.C. Huckett” (
Other material examined
12 males and females: Nearctic: Canada: Yukon Territory: 14 km WSW Burwash Flats, Firth River; USA: Alaska: Schrader L. Palaeartic: Kazakhstan: Dzungarian Alatau, Sarkand River; Russia: Altai Republic, Khakasiya, Tyva (BUIC,
Distribution
Nearctic: Canada (Yukon Territory); USA (Alaska). Palaearctic: Russia (Altai Mts, Khakasiya, Tyva), Kazakhstan.
DNA Barcode
BOLD BIN: BOLD:ADE2127. See Suppl. material
Remark
Only specimens from Russia were available for DNA barcoding (Fig.
Drymeia flavinervis
Pogonomyia flavinervis Malloch, 1915: 356.
Spilogaster nitens Stein, 1898: 199 [Junior primary homonym of Spilogaster nitens Macquart, 1855].
Pogonomyia flavipennis Stein, 1920: 21.
Type material examined
Pogonomyia flavinervis – Lectotype male labelled “
Other material examined
More than 100 males and females from: Nearctic: Canada: Alberta: Elkwater Park; Manitoba: Aweme, Bald Head Hills, 2 mi. E Douglas, 9 mi N. Forrest, Deloraine, Ninette, 30 mi N. Roblin, 3 mi. S. Shilo, 5 mi. SW Shilo, 2 mi. W Stockton, Teulon, Transcona, Turtle Mt., Virden; Nova Scotia: 4 km before Meat Cove (Victoria Co.); New Brunswick: Perth; Ontario: Bell’s Cor., Britannia, N. Burgess Twp., Burke Falls, Chatham, Constance L. (South March), Emo, Finland, 7 mi. E. Griffith, Hart L., Leamington, Metcalfe, Midland, Maynooth, March Twp., Marmora, Muskoka, Ojibway, Orillia, Ottawa, Point Pelee Ntl. Park, Pelee Is., Pinewood, Point Pelee, Rainy River, Rondeau Pr. Pk St. Lawrence Is. Nat. Park; Quebec: Alcove, Aylmer, Breackenridge; Gracefield, Laniel, Meach Lake, Mt. Auclair, Mt. Xalibu, Old Chelsea, Petit Mt St-Anne; Saskatchewan: Big River, Canora, Kenosee; USA: Maine: Tableland (Mt. Katahdin); Michigan: Isle Royale; Minnesota: Lake city; Virginia: Hawksbill (Shenandoah Ntl. Pk.) (BUIC, BIOUG,
Distribution
Nearctic: Canada (Alberta to New Brunswick), USA (South Dakota, Wisconsin, Illinois to Maine).
DNA Barcode
BOLD BIN: BOLD:ACA6790. See Suppl. material
Remark
DNA barcodes were available for material from Quebec and Ontario (Canada) with p-distances ranging from 0.0% to 0.16% (Fig.
Drymeia glacialis
Aspilia glacialis Rondani, 1866: 87.
Pogonomyia alpicola Rondani, 1871: 337. syn. nov.
Pogonomyia alpicola var. tundrica, Schnabl in
Type material examined
None.
Other material examined
More than 100 males and females: Nearctic: USA: Colorado: Cameron Pass, Cottonwood Pass (Chaffee Co.), Echo Lake (Mt. Evans), Floral Park, Independance Pass (Lake Co.), Loveland Pass, Nederland, Summit Lake (Mt. Evans); Wyoming: Delaey Creek Park, Snowy Range Mts., Togwotee Pass (Teton Co.). Palaearctic: Austria: Hohe Tauern Nat. Pk., Igls, Obergurgl; Italy: Karthaus; Mongolia: Ara-Khangaiskii aimak, 7 km SW Taryata; Russia: Altai Republic, Khakasiya, Tyva; Switzerland: Julierpass (BUIC,
Distribution
Nearctic: Canada (Alberta, Labrado,r and Newfoundland), USA (Rocky Mts down to New Mexico). Palaearctic: from Europe eastwards to the Far East of Russia.
DNA Barcode
BOLD BIN: BOLD:AAC1021 (BIN merge with D. quadrisetosa). See Suppl. material
Remarks
We were recently informed that the holotype of Aspilia glacialis Rondani, previously considered lost (
In the Nearctic region, females of this Holarctic species can be distinguished from those of D. quadrisetosa only by the slightly darker wing base. However, this colour character appears variable in the Palaearctic region where Russian material shows a darker wing base (congruent with Nearctic females) while females from a series we examined from Austria as well as the holotype of D. glacialis (A.C. Pont pers. comm.) display a pale wing base similar to that of D. quadrisetosa, a species known only from the Nearctic region.
DNA barcodes for D. glacialis (all from Russian specimens) were very similar to those of D. quadrisetosa (specimens from Russia and Canada), forming a cluster with p-distances ranging from 0.0% to 1.72% for BOLD:AAC1021 (Fig.
Drymeia groenlandica
Ophyra groenlandica Lundbeck, 1901: 281.
Type material examined
None.
Other material examined
More than 100 males and females: Nearctic: Canada: Manitoba: Churchill, Wapusk Ntl. Pk; Northwest Territories: Tuktoyaktuk; Nunavut: Arviat [formerly Eskimo point], Axel Heiberg Island, Aulavik Nat. Pk, Baker Lake, Coral Harbour (Southampton Island), Clyde (Baffin Island), Ellesmere Island, Eureka (Ellesmere Island), Forsheim peninsula (Ellesmere Island), Hazen Camp (Ellesmere Island), Kugluktuk [formerly Coppermine], Lake Hazen (Ellesmere Island), Masik River (Banks Island), Padlei, Tranquary Fiord (Ellesmere Island); Yukon Territory: North Fork Crossing mi 42 peel plt. Rd, Ogilvie Mountains. USA: Alaska: Schrader L. Greenland: Nedre Midsommer Sö, Zackenberg. Palaearctic: Russia: Chukotka: Pevek, Taymyr Peninsula: Ary-Mas cordon, Sakha Republik: Chokurdakh, Indigirka River (BIOUG, BUIC,
Male mid femur, anterior view (except G, H, L) A D. cantabrigensis B D. setibasis C D. vockerothi D D. profrontalis E D. glacialis F D. quadrisetosa G D. santamonicae, antero-dorsal view H D. amnicola, postero-dorsal view I D. similis J D. amnicola K D. hucketti L D. aldrichi, ventral view. Scale bars: 0.25 mm.
Distribution
Nearctic: Canada (Manitoba, Northwest Territories, Nunavut, Yukon Territory), USA (Alaska), Greenland. Palaearctic: Russia (Chukotka, Taymyr Peninsula, Sakha Republic).
DNA Barcode
BOLD BIN: BOLD:AAL9801. See Suppl. material
Remark
DNA barcodes for material from Canada, Greenland, and Russia (Fig.
Drymeia hucketti sp. nov.
Type material
Holotype
male labelled “Wood Mountain/ Sask 17.6.1955/ A.R. Brooks”; “HOLOTYPE/ Drymeia hucketti ♂/ Savage & Sorokina [red]” (
Other material examined
2 males: Nearctic: Canada: Alberta: Banff National Park; Saskatchewan: Grasslands National Park.
Etymology
The species name is a patronym in honour of Hugh C. Huckett, a major contributor to the study of Nearctic Muscidae.
Diagnosis
Small glossy black species with a long, strong prealar, lower margin of face projecting slightly beyond lower level of frons (Fig.
Description
Male. Body length: 4.6–6.6 mm; wing length 4.1–5.1 mm.
Head: Ground colour black; eye bare; fronto-orbital plate and parafacial silvery pruinose; face grey, gena and lower occiput grey pruinose; fronto-orbital plates touching in the middle; frons at narrowest point 2–3 × as wide as width of anterior ocellus; parafacial in lateral view with ventral margin broader than width of first flagellomere; lower margin of face projecting slightly beyond lower level of frons (Fig.
Thorax: Ground colour black; scutum, postpronotal lobe, notopleuron, postalar callus and pleuron subshiny and light grey dusted; anepimeron and katepimeron bare; notopleuron densely setulose; 2+3 dc; prealar long and strong.
Legs: Black; T1 with 1–3 pv; F2 with av row strong and regular, slightly longer than width of femur, covering apical 2/3 to 3/4 and with bristles curved on basal 1/2 (Fig.
Wing: Brown, darker near base; basicosta and tegula black; costal spinules weak and costal spine reduced; calypters with membrane and edges yellow.
Abdomen: Conical, ground colour black; lightly grey dusted and mostly shiny, tergites without distinct dark central vittae (Fig.
Terminalia: Fig.
Female. Body length: 5.7–6.5 mm; wing length: 4.6–5.0 mm (Fig.
Head: Frontal triangle undefined; ocellar triangle mostly glossy; frontal vitta black, deep brownish dusted; parafacial with large undusted shiny patch near base of antenna reaching up to or almost up to eye (Fig.
Thorax: As in male.
Legs (chaetotaxy described in full): T1 with 2 pv; F2 variable with 1–6 av (most specimens with 4 or 5), approximately as long as width of femur, and no pv; T2 with 2 or 3 ad, 3 or 4 pd, 2 pv; T3 with 3–5 av, 3–6 ad, 4 pd, and usually without pv (if present then very weak and short), apical pv distinct, at least 1/2 as long as apical av.
Wing: Veins yellow at least near base, membrane deep yellow near base, the remainder pale yellow to pale brown.
Abdomen: With little to no dusting, shiny.
Distribution
Nearctic: Canada (Alberta, British Columbia, Saskatchewan), USA (Montana).
DNA Barcode
BOLD BIN: BOLD:ACA9214. See Suppl. material
Remarks
The discovery of this species resulted from an exploration of all public Drymeia COI sequences found in BOLD (> 2800) which brought our attention to BOLD:ACA9214, a BIN including several well-preserved undetermined specimens of both sexes which turned out to be morphologically distinctive from any other species previously known to us. DNA barcodes for material from Canada and the United States (Fig.
Drymeia latifrons
Pogonomyia latifrons Malloch, 1918: 281.
Type material examined
Pogonomyia latifrons – Holotype female labelled “
Other material examined
None.
Distribution
Nearctic: USA (Colorado).
DNA Barcode
None available.
Remarks
The female holotype, with its unique combination of a broad frons, mid femur with weak av setae on apical 1/2, mid tibia with a single strong av, hind tibia with a strong apical pv, and wing membrane pale brown, is distinctive from any other specimen we have examined.
Drymeia minor
Pogonomyia minor Malloch, 1918: 280.
Type material examined
Pogonomyia minor – Holotype male labelled “Top of Las/ Vegas Range/VI.28.02 NM”; “HoloTYPE/6201 [red]”; “HoloTYPE/ Pogonomyia/ MINOR/ Mall. [red]” (
Other material examined
9 males, 3 females: Nearctic: USA: Colorado: Gunnison, Mt. Evans (
Distribution
DNA Barcode
BOLD BIN: BOLD:ADZ5293. See Suppl. material
Remarks
Based on the distribution of examined specimens (including the holotype of D. minor) as well as on the distribution of all public sequences for BOLD:ADZ5293 (D. minor) and BOLD:ACA9214 (D. hucketti sp. nov.), we suspect that previously published Canadian records of D. minor may actually belong to D. hucketti sp. nov. but additional data will be necessary for confirmation. Only specimens from Colorado (USA) were available for DNA barcoding (Fig.
Drymeia neoborealis
Aricia borealis Malloch, 1919: 64.
Helina neoborealis Snyder, 1949: 122 [replacement name for Aricia borealis Malloch, 1919].
Type material examined
Aricia borealis – Holotype male labelled “Bernard/ Harbour/ N.W.T./ July”; Canadian/ Arctic/ Expedition/ F. d. 1916”; TYPE/ H. borealis/ Mall./ No. 1176 [red]”/ “374” (
Other material examined
More than 50 males and females: Nearctic: Canada: Northwest Territories: Banks Island, Masik River, Victoria Island; Nunavut: Aulavik, Baker Lake, Cambridge bay, Char river nr Ranking Inlet, Chesterfield, Landing Lake (7.5 km NW of Rankin inlet). Palaearctic: Russia: Chukotka AO: Wrangel Island (BUIC,
Distribution
Nearctic: Canada (Northwest Territories, Nunavut); USA (Alaska, California, Colorado). Palaearctic: Russia (Wrangel Island).
DNA Barcode
BOLD BIN: BOLD:ACA8935. See Suppl. material
Remarks
Drymeia ponti sp. nov.
Type material
Holotype
male labelled “W. Side Cortes Pass/ 1100' Mexico, Mexico/ 13-VIII-1954/ J.G. Chillcott”; “HOLOTYPE/ Drymeia ponti ♂/ Savage & Sorokina [red]” (
Etymology
Diagnosis
Small dark species with strong prealar, 2+3 dc and strong costal spine. This species is similar to Drymeia aterrima (Wulp, 1896), especially in the dark male calypter, but can be distinguished from it in the male by a broad frontal vitta (Fig.
Description
Male. Body length: 4.1–5.7 mm; wing length: 3.8–5.2 mm.
Head: Ground colour black; eye bare; fronto-orbital plate and parafacial dark brown pruinose; face black, gena and lower occiput dark brown pruinose; frons at narrowest point approximately 1.5 × width of ocellar triangle with black frontal vitta exposed (Fig.
Thorax: Ground colour black; scutum, postpronotum, notopleuron, postalar callus and scutellum with dense brown dusting; pleuron dark brown, slightly shiny with brown dusting; katepisternum with small anterior undusted glossy patch, meron entirely dusted; anepimeron and katepimeron bare; notopleuron setulose; acr 0+1; 2+3 dc; prealar long and strong, as long as second notopleural.
Legs: Black; T1 with 2 pv on apical 1/2; F2 straight, with matching rows of long strong av and pv on apical 2/3, these much longer than width of femur (Fig.
Wing: Brown, darker at base with dark brown veins; basicosta and tegula black; costal spinules strong, with costal spine 2 × as long as costal spinules; calypters with membrane and edges dark brown.
Abdomen: Conical; ground colour black; brown dusted, subshiny with no median vittae; sternite I bare; sternite V as in Fig.
Terminalia: Fig.
Female. Body length: 4.5–6.5 mm; wing length: 4.0–5.5 mm. Differs from the male as follows:
Head: Frontal triangle undefined; frontal vitta black; parafacial mostly dusted, with a small narrow shiny patch near base of antenna (Fig.
Thorax: As in male.
Legs
(chaetotaxy described in full): T1 with 1 or 2 pv; fore tarsomere 5 distinctively flattened (Fig.
Wing: Light brown, darker near base; with membrane and edges dark yellow.
Abdomen: as in male.
Distribution
Neotropical: Mexico (Mexico).
DNA Barcode
None available.
Drymeia pribilofensis
Eupogonomyia pribilofensis Malloch, 1921: 179.
Pogonomyia inaequalis Malloch, 1922: 81.
Type material examined
Eupogonomyia pribilofensis – Holotype male labelled “St. Paul Isd./ Alaska/ VI-21-20”; “Presented by/ G.D. Hanna/ Collector”/ “HOLOTYPE/ pribilifensis [red]”; “Eupogonomyia/ pribilofensis/ Mall. Type” (
Other material examined
More than 400 males and females from: Nearctic: Canada: Manitoba: Churchill; Northwest Territories: Aklavik, Aulavik Nt. Pk. (Banks Island), Kidluit Bay, Mould Bay, Tuktoyaktuk; Nunavut: Arviat [formerly Eskimo point], Bathurst Inlet, Cambridge Bay, Chesterfield, Coral Harbour, Frobisher Bay, Kugluktuk [formerly Coppermine], Landing Lake (7.5 km NW of Rankin Inlet), Naujaat [formerly Repulse Bay], Padlei, Williamson Lake; Quebec: Inukjuak [formerly Port Harrison], Kangirsuk [formerly Payne Bay], Sugluk; Yukon Territory: Herschel Island. Palaearctic: Russia: Taymyr Peninsula: Ary-Mas cordon, 90 km NW Khatanga, Dixon; Yamalo-Nenez AO; Chukotka AO: Wrangel Island; Sakha Republic: Chokurdakh, Indigirka River (BUIC,
Distribution
DNA Barcode
BOLD BIN: BOLD:AAD7664 (BIN merge with several other species, see Fig.
Remark
Males of this species can be readily identified based on the combination of a reduced prealar, strongly projecting face (Fig.
Drymeia profrontalis
Pogonomyia profrontalis Huckett, 1966: 293.
Type material examined
Pogonomyia profrontalis – Holotype male labelled “Big Spring/ Shasta Country/ Calif. v-23:41”; “E. G. Linsley/ Collector”; progonomyia/ profrontalis/ n. sp./ holotype [red]”; “California Academy/ of Sciences/ Type No. 10150” (
Other material examined
5 males, 3 females: Nearctic: USA: California: Buck’s Lake, Wright’s Lake (
Distribution
Nearctic: USA (California).
Drymeia quadrisetosa
Pogonomyia quadrisetosa
Pogonomyia amurensis Lavčiev, 1971: 220
Type material examined
None.
Other material examined
More than 300 males and females from: Nearctic: Canada: Manitoba: Churchill, Fort Churchill; Labrador: Cutthroat Harb., Hebron, Nutak; Northwest Territories: Muskox L., Salmita Mines, Tuktoyaktuk, 21 mi E. Tuktoyaktuk; Wholdaia Lake; Victoria Island; Nunavut: Baker Lake, Bathurst Inlet, Cambridge Bay, Chesterfield, Kugluktuk [formerly Coppermine], Landing Lake (7.5 km NW of Rankin Inlet), Padlei; Quebec: Kangirsuk [formerly Payne Bay], Sugluk; Yukon Territory: British Mts, 17 km WNW Burnash Flats, km 159 Dempster Highway, Firth River, Herschel Island; USA: Alaska: Cape Thompson, Isabel Pass, Schrader L., Umiat, Unalakleet [Noluk 68N, 160W]. Palaearctic: Russia: Taymyr Peninsula: 90 km NW Khatanga; Yamalo-Nenez AO: 73 km NE Labytnangy; Chukotka AO: Pevek; Republic of Buryatia: Baisa; Sakha Republic: 100 km NW Oymyakon (BUIC,
Distribution
Nearctic: Canada (Manitoba, Northwest Territories, Nunavut, Quebec, Yukon Territory; USA (Alaska). Palaearctic: Russia (Siberia).
DNA Barcode
BOLD BIN: BOLD:AAC1021 (BIN merge with D. glacialis). See Suppl. material
Remarks
See comments about DNA barcodes under D. glacialis.
Drymeia santamonicae
Pogonomyia santamonicae Huckett, 1966: 294.
Type material examined
Pogonomyia santamonicae – Holotype male labelled “StaMonica Mts./ L.A. Co. Cal./ VII-3-50”; “Pogonomyia/ santamonicae/ n. sp./ holotype [red]”; “California Academy/ of Sciences/ Type No. 10151” (
Other material examined
7 males, 1 female: Nearctic: USA: California: Cleaveland Nat. Forest, Cuyamaca L., Laguna Jct. (
Distribution
Nearctic: USA (California).
DNA Barcode
None available.
Drymeia segnis
Aricia segnis Holmgren, 1883: 169.
Pogonomyioides atrata Malloch, 1919: 67.
Type material examined
Pogonomyioides atrata Malloch. Holotype female (with puparium) labelled “Bernard/ Harbour/ N.W.T./July 7. [vertical]”; “Canadian/ Arctic/ Expedition/ F.J. 1915”; “1215”/ “Type/ No. 1180 [red]”; “SLIDE Coll./ A 162 [blue]” [head mounted on slide] (
Other material examined
More than 500 males and females: Nearctic: Canada: Alberta: Eisenhower Jct., Snow Creek Pass (Banff N. P.); Northwest Territories: Holman, Banks Island (Aulavik, Masik river), Sachs Harbour, Tuktoyaktuk; Victoria island; Nunavut: Alex Fiord, Arviat [formerly Eskimo point], Axel Heiberg Island, Baker Lake, Cambridge Bay, Chesterfield, Clyde, Coral Harbour, Devon Island, Ellesmere island, Eureka, Hazen Camp, Kugluktuk, Landing Lake (7.5 km NW of Rankin inlet), Meliadine river, Naujaat [formerly Repulse Bay], Taloyoak [formerly Spence Bay], Tranquary Fjord; Quebec: Inukjuak [formerly Port Harrison]; Yukon Territory: British Mts, Firth River, Herschel Island, Richardson Mts. USA: Alaska: Upper Colville River; Colorado: Mt. Evans. Greenland: Nedre Midsommer Sö. Palaearctic: Russia: Taymyr Peninsula: 90 km NW Khatanga, Ary-Mas cordon, Dixon; Chukotka AO: Wrangel Island; Sakha Republic: 19 km SE Kyusyur, (BUIC,
Distribution
Nearctic: Canada (Alberta, Manitoba, Northwest Territories, Nunavut, Quebec, Yukon Territory), USA (Alaska, Colorado), Greenland. Palaearctic: Russia.
DNA Barcode
BOLD BIN: BOLD:AAD7664 (BIN merge with several other species, see Fig.
Remark
DNA barcodes for D. segnis material from Canada and Greenland were very similar to several of the seven species found in the BOLD:AAD7664 BIN merge (Fig.
Drymeia setibasis
Eupogonomyia setibasis Huckett, 1965: 301.
Thrichopticoides gymnophthalma sibirica Lavčiev, 1971: 220.
Type material examined
Eupogonomyia setibasis – Holotype male labelled “Lady Melville L./ 93°15'W, 69°25'N/ NWT 3.VII 1951”; “Type [red]”; “HOLOTYPE/ CNCNo. 8368 [red]”; “Eupogonomyia/ setibasis Huck./ det. H.C. Huckett” (
Other material examined
More than 50 males and females from Nearctic: Canada: Northwest Territories: Tuktoyaktuk; Nunavut: Arviat [formerly Eskimo point], Cambridge Bay, Landing Lake (7.5 km NW of Rankin inlet), Taloyoak [formerly Spence Bay]; Yukon Territory: 17 km WNW Burwash Flats. Palaearctic: Russia: Taymyr Peninsula: Ary-Mas cordon, 12.5 km S. Dikson settlement, 90 km NW Khatanga; Altai Republic; Chukotka AO: Wrangel Island (BUIC,
Distribution
Nearctic: Canada (Northwest Territories, Nunavut, Yukon Territory), USA (Alaska). Palaearctic: Armenia, Georgia, Kyrgyzstan, Russia.
DNA Barcode
BOLD BIN: BOLD:AAD7664 (BIN merge with several other species, see Fig.
Remarks
See comments under D. segnis and D. cantabrigensis.
Drymeia similis
Pogonomyia similis Malloch, 1918: 279.
Type material examined
Pogonomyia similis Malloch – Holotype male labelled “Top of Las/ Vegas Range/VI.28.02 NM”; “HoloTYPE/6200 [red]”; “HoloTYPE/ Pogonomyia/ MINOR/ Mall. [red]” (
Other material examined
Nearctic: Canada: Alberta: Acme, Banff, 20 mi W Calgary, Drumheller, Eisenhower Jct., Elkwater Lake, Frank, Hinton, Lethbridge, 15 km east Morley, Waterton, British Columbia: Atlin, Cathedral Mtn, Clinton, Crowsnest, Liard Hot Spg., Vernon, Victoria; Manitoba: Carberry, Gimli, Husavick, Pierson, Reynolds, 30 mi N Roblin, Taulon[a], Virden; Northwest Territories: Fort Liard, Hay River; Ontario: Moosonee, Thor Lake; Quebec: Mistassini, Rupert House; Saskatchewan: Big River, Cypress Hills, Kenosee, Melfort, Prince Albert; Yukon Territory: Dawson. USA: Colorado: Chicago Cr., Estes Park, Gilpin, Idaho Springs, Jefferson, Loveland Pass, Mt. Evans (Doolitle Ranch, Echo Lake), Nederland, Niwot Ridge; Utah: Daniels Pass; Wyoming: Togwotee Pass, Union Pass Road. Greenland: Nedre Midsommer Sö. (
Distribution
Nearctic: Canada (Yukon Territory and British Columbia to Newfoundland and Labrador), USA (Alaska to New Mexico), Greenland.
DNA Barcode
BOLD BIN: BOLD:AAG1776. See Suppl. material
Remark
All three specimens of D. similis in our DNA barcoding data set were from Saskatchewan (Canada) with p-distances ranging from 0.0% to 0.16% (Fig.
Drymeia spinitarsis
Pogonomyia spinitarsis Aldrich, 1918: 184.
Drymeia longiseta Sorokina & Pont, 2015: 181. syn. nov.
Type material examined
Drymeia longiseta Sorokina & Pont. Holotype male labelled “Russia, Republic Altai/ Kosh-Agash area, 7 km NW/ Kuray, 2251 m, Kurayskiy/ mt. ridge, 50°17'N, 87°51'E/ Coll. V. Sorokina, 17.07.2013”; “Holotype/ Drymeia longiseta ♂/ Sorokina & Pont sp.n. [red]” (
Other material examined
Over 400 males and females from: Nearctic: USA: Colorado: Corona Pass, Cottonwood Pass, Estes Park, Loveland Pass, Mt. Evans, Nederland, Niwot Ridge, Summit Lake (Mt. Evans) (
Distribution
Nearctic: USA (Colorado). Palaearctic: Russia (Altai-Sayan region).
DNA Barcode
BOLD BIN: BOLD:ACT4320. See Suppl. material
Remarks
The comparison of type material from D. longiseta Sorokina and Pont from Russia with a series of high elevation specimens from Colorado, USA, housed in the
Drymeia vockerothi sp. nov.
Type material
Holotype
male labelled “Rigaud, QUE./ 11.VI. 1981/ J.R. Vockeroth”; “Summit of/ Mt. Rigaud”; “
Etymology
The species name is a patronym in honour of famous dipterologist John R. Vokeroth (Canada), who collected the holotype and several paratypes and designated these specimens as a “new species near D. neoborealis”.
Diagnosis
Large glossy black species with short proboscis, very narrow parafacial in lateral view, arista almost bare, prealar weak or absent, 2+3 or 4 dc and well developed presutural acr. Males with narrow frons (Fig.
Male. Body length: 7.2–9.5 mm; wing length: 6.0–6.7 mm (Fig.
Head: Ground colour black; eye bare; fronto-orbital plate and parafacial yellowish silver pruinose; face grey, gena and lower occiput grey pruinose; frons at narrowest point 1.5–2.5 × width of anterior ocellus with fronto-orbital plates touching or nearly touching (Fig.
Thorax: Ground colour black; scutum, postpronotum, notopleuron, postalar callus, pleuron and scutellum black and shiny, mostly undusted; large undusted glossy patches on anterior part of katepisternum and ventral area of meron; anepimeron and katepimeron bare; notopleuron densely setulose; acr 2 or 3+4 or 5, stronger than ground setulae and with preapical series at least as long as preapical dc; 2+4 dc; prealar weak or absent, always much shorter than 2nd notopleural.
Legs: Black; T1 with 1 or 2 pv; F2 straight, slightly flattened anterodorsally, with 3 or 4 long delicate upcurved av on basal 1/3 and short setae on remaining av surface (Fig.
Wing: Light brownish, yellow at base; basicosta and tegula black; costal spinules weak and costal spine reduced; calypters with membrane and edges yellow.
Abdomen: Conical, ground colour black; densely grey dusted with narrow black median vittae on tergites II–V; sternite I bare; sternite V as in Fig.
Terminalia: Fig.
Female. Body length: 7.5–9.0 mm; wing length: 6.2–6.8 mm (Fig.
Differs from the male as follows:
Head: Ground colour black with grey dusting; frontal vitta brown with grey dusting; frontal triangle indistinct, ocellar triangle undusted and shiny; parafacial entirely dusted, without shiny patch near base of antenna; frons at midpoint 0.36 × as wide as head and approximately 0.85 × as long as wide; fronto-orbital plate wide, at midpoint 0.4 × as wide as frontal vitta; seven medioclinate frontal setae (including several interstitials), three orbital setae, the upper two reclinate and lateroclinate, the lower one proclinate.
Thorax: Ground colour as in male but with light grey dusting (Fig.
Legs (chaetotaxy described in full): T1 with 1 or 2 pv; F2 not flattened anteroventrally, with 2 or 3 short av, 2 or 3 short v on basal 1/3, and with a few short fine pv on apical 1/3; T2 with 1 or 2 ad, 3–5 pd and 1 or 2 pv; F3 with av row stronger on apical 1/2, without pv; T3 with 3 or 4 short av, 3 or 4 ad, and 3–6 pd, apical pv absent.
Wing: As in male.
Abdomen: Without distinct median vittae.
Distribution
Nearctic: Canada (New Brunswick, Ontario, Quebec).
DNA Barcode
None available.
Drymeia woodorum sp. nov.
Type material
Holotype
male labelled “MI. 87, Y.T./ Depmster Hwy./ 4–8.VIII.1973/ G. & D.M Wood”; “HOLOTYPE/ Drymeia woodorum ♂/ Savage & Sorokina [red]” (
Etymology
The species name is a patronym in honour of the late Canadian dipterologist D. Monty Wood and his wife Grace Wood (Canada), who collected the holotype.
Diagnosis
Small grey species with short proboscis, narrow frons and narrow parafacial in lateral view (Fig.
Description
Male. Body length: 5.5 mm wing length: 4.5 mm.
Head: Ground colour black; eye bare; ocellar triangle, fronto-orbital plate and parafacial silvery pruinose; face grey, gena and lower occiput light grey pruinose; fronto-orbital plates touching in the middle; frons at narrowest point as wide as width of anterior ocellus; parafacial very narrow in lateral view, < 1/2 width of first flagellomere along full length (Fig.
Thorax: Ground colour black; scutum without distinct vittae; postpronotum, notopleuron and postalar callus with dense brown dusting; pleuron brownish grey dusted; undusted glossy patches on most anterior part of katepisternum and on a small ventral area of meron; anepimeron and katepimeron bare; notopleuron densely setulose; acr 2 or 3 + 4 or 5 (+1 prescutellar), stronger and longer than ground setulae (and with preapical series at least as long as preapical dc); 2 or 3+4 dc; prealar absent.
Legs: Black; fore coxa with patch of four or five very long, strong apical bristles, at least as long as length of coxa; T1 with 1 or 2 pv; F2 with a row of short av on apical 1/3, 5 or 6 a on basal 1/2, these longer than width of femur, 3 preapical pd-p, and dense fine pv over most of the surface; T2 without av, with a row of ad on apical 2/3, these longer near apex, 4 pd and 2 pv; F3 with av row stronger on apical 1/3, without pv except for a few hairs near base; T3 with 3 av, 3 long ad, a row each of short erect uneven a and ad, 2 long and 2 or 3 short pd, ventral apical process very short but distinct, apical pv absent (Fig.
Wing: Mostly dark brown; basicosta and tegula black; costa with short weak spinules and costal spine reduced; calypters with membrane and edges deep yellow.
Abdomen: Conical, ground colour black; densely grey dusted with a black median vitta on tergites II–V; sternite I bare; sternite V as in Fig.
Terminalia: Fig.
Female. Body length: 4.9–5.5 mm wing length: 4.3–4.7 mm (Fig.
Head: Ground colour grey with grey dusting; frontal vitta black with brownish dust; frontal triangle indistinct, ocellar triangle covered with heavy grey or brown dust; parafacial completely dusted but with large area near base of antenna appearing velvety brown in anterior and lateral views (Fig.
Thorax: Scutum with whitish grey dust and, when viewed from the front, with 3 brown vittae with diffuse margins (Fig.
Legs (chaetotaxy described in full): Fore coxa without patch of long strong bristles; T1 with 1 or 2 pv; F2 with a row of short av, no longer than width of femur and a row of delicate pv-v, no longer than diameter of femur; T2 usually with a row of 5–7 irregular pd, (these sometimes reduced) and 1 pv; F3 with av row stronger on apical 1/3, without pv except for a few hairs near base, T3 with 2–3 av, 5–6 short irregular ad and 5–6 pd, apical pv usually absent but if visible, then no longer than 1/2 the length of apical av.
Wing: Clear with slight pale brown to pale yellow tinge, veins pale brown; calypters whitish.
Abdomen: Densely whitish grey dusted with slight brownish tinge along posterior margins of tergites II–IV.
Distribution
Nearctic: Canada (Yukon Territory).
DNA Barcode
None available.
Remarks
See notes on problematic taxa.
Notes on problematic North American taxa
Neighbour-joining tree based on p-distance of all COI sequences (> 550 bp) for which specimens were examined. Each terminal includes the species name, BOLD process ID number, country of collection, and Barcode Index Number. Red text indicates the presence of more than one named species grouped under a single BIN (BIN merge).
Drymeia vockerothi sp. nov. is another species with a short prealar, short proboscis and proclinate orbital setae but it is much larger than the other taxa discussed above and does not fit the features listed in the key or the distribution of any Trichopticoides spp. mentioned in
Morphology
Three new synonymies are established here (the recognition of D. longiseta Sorokina & Pont, 2015 as a junior synonym of D. spinitarsis (Aldrich, 1918), of D. rivalis (Huckett, 1966) as a junior synonym of D. amnicola (Huckett, 1966), and of Drymeia alpicola (Rondani, 1871) as a junior synonym of Drymeia glacialis (Rondani, 1866)). These synonymies are based exclusively on morphology and details are provided in the checklist section of this work.
Most species of Drymeia show distinctive and discrete morphological differences between species (at least in the males). However, the four species described in
DNA barcoding
A total of 175 sequences were included in the DNA barcode data set (Suppl. material
Eighteen named species and four unknown taxa matched a single BIN. These numbers were compiled after we re-examined material (all paratypes) from BOLD:ACT1697 (males and females assigned to D. grandis Sorokina & Pont, 2015) and BOLD:ACT1698 (a mixture of males D. cilitarsis Sorokina & Pont, 2015, and females originally assigned to D. grandis). The female of D. cilitarsis is currently undescribed and the re-examination of specimens from these two BINs led us to the conclusion that the paratype series of D. grandis is mixed, containing properly identified females as well as some belonging to D. cilitarsis (both taxa have similar distributions). The description of the female of D. cilitarsis, a Palaearctic species, is outside the scope of the present work and will be completed in a separate contribution.
Following the examination of material from the four BINs containing specimens we could not assign to known species based on morphology we described one new species, Drymeia hucketti sp. nov. (BOLD:ACA9214). Due to the very poor condition of available specimens (BOLD:ACA9236), combined with a lack of males for BOLD:ACZ1539 we chose to leave these two clusters unassigned as Drymeia sp. 1 and Drymeia sp. 2, respectively (Fig.
The four specimens (two from each sex) available for BOLD:ACY7523 (Drymeia sp. nr. aldrichi), were most similar to D. aldrichi Malloch, 1918 but differed slightly from material examined (holotype male + two females from same series) in the following features: females with very short av and pv near the base of the mid femur and with tarsomere 5 not flattened, males with projection of frons slightly more pronounced and with slightly longer pv on the apical 1/2 of the mid femur. Since no DNA barcodes were available for material matching the morphology of specimens of D. aldrichi examined here, we chose to leave this cluster unresolved as we are unable at present time to determine if the material from BOLD:ACY7523 belongs to a new distinct species or if D. aldrichi is more variable morphologically than currently understood.
While the congruence between BIN assignment and morphology was good for 71% (22/31) of species studied here (including the three unnamed taxa), a significant proportion of species (9/31) presented no or very low interspecific distance to other morphologically distinct taxa. These taxa were grouped under two BIN “merges” including two (BOLD:AAC1021) and seven (BOLD:AAD7664) species (Fig.
Low interspecific distance between sequences from Canadian specimens of D. pribilofensis and D. segnis had previously been flagged by
Male specimens were included in our dataset for all species involved in the two BIN merges except D. puchokana; we therefore exclude misidentification as a possible explanation for the mismatches between BIN assignment and morphology reported here. Low genetic divergence in COI sequences between closely related species has been reported in the literature for different groups of animals (e.g.,
Conclusions
We found both advantages and limitations to using DNA barcodes and BIN assignments for the identification of Drymeia species. They allowed us to resolve the issue of male/female associations in the Palaearctic species D. grandis and D. cilitarsis, to discover one new species, D. hucketti sp. nov., and to focus our attention on three potentially new taxa (Drymeia sp. 1, D. sp. 2, D. sp. nr. aldrichi). BINs were also a good match for 22 of the 31 species included in our data set, regardless of geographical distance. However, the nine remaining species could not be distinguished from at least one other species; we therefore caution against using BIN assignments based on COI DNA barcoding as the only determination tool for Drymeia material without prior knowledge of its limitations for certain species groups. This is especially important for the Nearctic fauna where only 65% (11/17) of the species represented in our DNA barcode library (including the three unnamed taxa) have a match between BIN and morphology while the remaining six show no or very low sequence divergence with one or more morphologically distinct species.
Considering the taxonomic changes presented here, the North American fauna of Drymeia now includes 24 species; 22 are found in the Nearctic region, D. ponti sp. nov. is recorded only from Neotropical Mexico, and D. aterrima has been recorded from both the Nearctic and Neotropical regions. Furthermore, the synonymy of D. longiseta with D. spinitarsis now brings the number of Holarctic species to ten. Whereas the present study provides the first comprehensive taxonomic treatment of Drymeia for North America, ten of the 24 named species of Drymeia are still missing DNA barcodes and some additional work remains. The most important gap in knowledge concerns the fauna of the southwestern United States and Mexico and the collection of additional specimens from those areas are likely to result in further changes to the taxonomy of the genus.
Acknowledgements
We thank the following people and institutions for specimen loans: Stéphanie Boucher (LEM), Roberta Brett (
We are grateful to Jeffrey Skevingon (
Financial support was provided by an NSERC Discovery grant (Canada) to J. Savage, Bishop’s University, a CanaColl grant (CanaColl foundation) to V.S. Sorokina and the work of V.S. Sorokina was carried out within the framework of the State Research Program FWSG-2021-0004. The authors have declared that no competing interests exist.
References
- Agassiz L (1847) Nomenclatoris zoologici index universalis, continens nomina systematic classium, ordinum, familiarum et generum animalium omnium, tam viventium quam fossilium, secundum ordinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nec non variis adnotationibus et emendationibus (1846). Jent and Gassmann. Soloduri [= Solothurn, Switzerland], 393 pp. https://doi.org/10.5962/bhl.title.1819
- Aldrich JM (1918) The Anthomyid genus Pogonomyia (Dip.). Entomological news 29(5): 179–185.
- Becker T, Dziedzicki H, Schnabl J, Villeneuve J (1915) Résultats scientifiques de l’Expédition des frères Kuznecov (Kouznetzov) à l’Oural Arctique en 1909, sous la direction de H. Backlund. Diptera. Zapiski Imperatorskoĭ Akademii Nauk (VIII), Fiziko–Matematicheskoe Otdeleniye 28(7): 1–67.
- Carvalho CJB de, Couri MS, Pont AC, Pamplona D, Lopes SM (2005) A catalogue of the Muscidae (Diptera) of the Neotropical Region. Zootaxa 860: 1–282. https://doi.org/10.11646/zootaxa.860.1.1
- Fan Zide (2008) Fauna Sinica. Insecta (Vol. 49). Diptera Muscidae (1). Science Press, Beijing, [xvi +] 1186 pp.
- Foottit RG, Maw HEL, Havill NP, Ahern RG, Montgomery ME (2009) DNA barcodes to identify species and explore diversity in the Adelgidae (Insecta: Hemiptera: Aphidoidea). Molecular Ecology Resources 9: 188–195. https://doi.org/10.1111/j.1755-0998.2009.02644.x
- Funk DJ, Omland KE (2003) Species-Level Paraphyly and Polyphyly: Frequency, Causes, and Consequences, with Insights from Animal Mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics 34(1): 397–423. https://doi.org/10.1146/annurev.ecolsys.34.011802.132421
- Gibbs J (2018) DNA barcoding a nightmare taxon: assessing barcode index numbers and barcode gaps for sweat bees. Genome 61(1): 21–31. https://doi.org/10.1139/gen-2017-0096
- Gistl J (1848) Naturgeschichte des Thierreichs für höhere Schulen. Hoffmann’sche Verlags-Buchhandlung, Stuttgart, 216 pp.
- Gregor F, Rozkošný R, Barták M, Vaňhara J (2016) Manual of Central Europe Muscidae (Diptera). Schweizerbart Science Publishers (Nägele u. Obermiller), Zoologica 162, Stuttgart, 219 pp.
- Hajibabaei M, deWaard JR, Ivanova NV, Ratnasingham S, Dooph RT, Kirk SL, Mackie PM, Hebert PDN (2005) Critical factors for assembling a high volume of DNA barcodes. Philosophical Transactions of the Royal Society B 360: 1959–1967. https://doi.org/10.1098/rstb.2005.1727
- Hebert PDN, Penton EH, Burns JM, Janzen DH, Hallwachs W (2004) Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America 101: 14812–14817. https://doi.org/10.1073/pnas.0406166101
- Hennig W (1962a) 63b. Muscidae. [Part, Lieferung 225.]. In: Lindner E (Ed.) Die Fliegen der palaearktischen Region. Schweizerbart, Stuttgart, 625–672.
- Hennig W (1962b) 63b. Muscidae. [Part, Lieferung 227.]. In: Lindner E (Ed.) Die Fliegen der palaearktischen Region. Schweizerbart, Stuttgart, 673–720.
- Holmgren AE (1883) Insecta a viris doctissimis Nordenskiöld illum ducem sequentibus in Insulis Waigatsch et Novaja Semlia anno 1875 collecta. Diptera. Entomologisk Tidskrift 4: 162–190.
- Huckett HC (1965a) The Muscidae of Northern Canada, Alaska and Greenland (Diptera). Memoirs of the Entomological Society of Canada 42: 1–370. https://doi.org/10.4039/entm9742fv
- Huckett HC (1965b) Family Muscidae. In: Stone A, Sabrosky CW, Wirth WW, Foote RH, Coulson JR (Eds) A Catalog of the Diptera of America North of Mexico. Agricultural Handbook of the United States Department of Agriculture 276: 869–915.
- Huckett HC (1966) New species of Anthomyiidae and Muscidae from California (Diptera). Proceedings of the California Academy of Sciences 34(3): 235–306.
- Huckett HC (1975) The Muscidae of California exclusive of subfamilies Muscinae and Stomoxyinae. Bulletin of the California Insect Survey 18: 1–148.
- Huckett HC, Vockeroth JR (1987) Muscidae. In: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM (Eds) Manual of Nearctic Diptera Volume 2. Agriculture Canada Monograph 28: 1115–1131.
- Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution 35: 1547–1549. https://doi.org/10.1093/molbev/msy096
- Lavčiev VI (1971) Faunistische und taxonomische Bemerkungen über einige Musciden (Diptera, Muscidae) aus UdSSR nebst Beschreibungen der neuen Arten. Entomologicheskoe Obozrenie 50: 215–226. [In Russian, with German summary] [English translation in Entomological Review 50: 125–131.]
- Lundbeck W (1901) Diptera groenlandica. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening (6) 2(1900): 281–316.
- Malloch JR (1915) Some additional records of Chironomidae for Illinois and notes on other Illiois Diptera. Ill. Bulletin of the Illinois State Laboratory of Natural History (1918) 11: 305–363. https://doi.org/10.21900/j.inhs.v11.357
- Malloch JR (1918) Diptera from the south-western United States. Part IV. Anthomyiidae. Transactions of the American Entomological Society 44: 263–319.
- Malloch JR (1919) The Diptera collected by the Canadian Expedition, 1913–1918 (Excluding the Tipulidae and Culicidae). In: Anderson RM (Ed.) Report of the Canadian Arctic Expedition 1913–18 (Vol. 3). Insects. Part C. Diptera. Printer to the King’s Most Excellent Majesty Ottawa, Thomas Mulvey, Ontario, 34–90.
- Malloch JR (1921) Dipterous insects of the family Anthomyiidae from from the Pribilof Islands, Alaska. Proceedings of the California Academy of Sciences (4) 11: 178–182.
- Malloch JR (1922) A new European anthomyiid (Diptera). Notulae Entomologicae 2: 81–82.
- McAlpine JF (1981) Morphology and terminology – Adults. In: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM (Eds) Manual of Nearctic Diptera Vol. 1. Agriculture Canada Monograph 27: 9–63.
- Meigen JW (1826) Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten. Bei Friedrich Wilhelm Forstmann: Gedrukt bei Beaufort Sohn Hamm 5: 1–412.
- Michelsen V (2011) A new European species of Drymeia Meigen (Diptera: Muscidae) near D. brumalis (Rondani), with observations of a unique prestomal teeth structure. Zootaxa 3023: 51–60. https://doi.org/10.11646/zootaxa.3023.1.3
- Michelsen V (2015) Anthomyiidae, Fanniidae, Muscidae and Scathophagidae (The Muscidae family group). In: Böcher J, Kristensen NP†, Pape T, Vilhelmsen L (Eds) The Greenland Entomofauna 17, Diptera. Brill, Leiden and Boston, 635–657.
- Morrone JJ (2014) Biogeographical regionalization of the Neotropical region. Zootaxa 3782: 001–110. https://doi.org/10.11646/zootaxa.3782.1.1
- Nihei SS, Carvalho de CJB de (2004) Lectotype designation and redescription of Drymeia aterrima (Wulp, 1896) (Diptera: Muscidae). Zootaxa 612: 1–6. https://doi.org/10.11646/zootaxa.612.1.1
- Pont AC (1981) Himalayan Muscidae. III. The genus Pogonomyia Rondani (Diptera). Spixiana 4: 121–142.
- Pont AC (1986) Family Muscidae. In: Soós Á, Papp L (Eds) Catalogue of Palaearctic Diptera 11, Scathophagidae – Hypodermatidae. Akadémiai Kiadó, Budapest, 57–215.
- Ratnasingham S, Hebert PDN (2013) A DNA-Based Registry for All Animal Species: The Barcode Index Number (BIN) System. PLoS ONE 8(7): e66213. https://doi.org/10.1371/journal.pone.0066213
- Renaud AK, Savage J, Adamowicz SJ (2012) DNA barcoding of Northern Nearctic Muscidae (Diptera) reveals high correspondence between morphological and molecular species limits. BMC Ecology 12: e24. https://doi.org/10.1186/1472-6785-12-24
- Ringdahl O (1931) Einige Mitteilungen über lappländische Dipteren. Entomologisk Tidskrift 52: 171–174.
- Rondani C (1866) Anthomyinae Italicae collectae distinctae et in ordinem dispositae [a prof. Camillo Rondani]. Atti della Società Italiana di Scienza Naturali 9: 68–217.
- Rondani C (1871) Diptera Italica non vel minus cognita descripta aut annotate. Fasc. IV. Addenda Anthomyinis Prodr. (Vol VI). Bollettino della Societa entomologica inaliana 2(1870): 317–338.
- Savage J, Wheeler TA (2004) Phylogeny of the Azeliini (Diptera: Muscidae). Studia Dipterologica 11(1): 259–299.
- Schnabl J, Dziedzicki H (1911) Die Anthomyiden. Nova Acta Academiae Caesarea Leopoldina-Carolinae Germanicum Naturae Curiosorum 95: 55–358.
- Séguy E (1923) Diptères Anthomyides. Faune de France 6: [xi +] 393 pp.
- Shinonaga S (2007) Muscidae from Pakistan (Diptera). Japanese Journal of Systematic Entomology 13: 267–311.
- Shinonaga S, Singh MM (1994) Muscidae of Nepal (Diptera). I. Muscinae, Stomoxyinae and Phanoniinae. Japanese Journal of Sanitary Zoology 45 (Suppl.): 99–177. https://doi.org/10.7601/mez.45.99
- Skidmore P (1985) The biology of the Muscidae of the world. Series Entomologica 29: е550.
- Snyder FM (1949b) Nearctic Helina Robineau-Desvoidy (Diptera: Muscidae). Bulletin of the American Museum of Natural History 94: 107–160.
- Sorokina VS (2012) The Muscidae (Diptera) of the Russian tundra zones. Report 1. Caucasian Entomological Bulletin 8(2): 328–332. [In Russian, with English summary] https://doi.org/10.23885/1814-3326-2012-8-2-328-332
- Sorokina VS, Pont AC (2010) An annotated catalogue of the Muscidae (Diptera) of Siberia. Zootaxa 2597: 1–87. https://doi.org/10.11646/zootaxa.2597.1.1
- Sorokina VS, Pont AC (2015) A review of the genus Drymeia Meigen, 1826 (Diptera: Muscidae) in Russia. Zootaxa 4000(2): 151–206. https://doi.org/10.11646/zootaxa.4000.2.1
- Stein P (1898) Nordamerikanische Anthomyiden. Beitrag zur Dipterenfauna der Vereinigten Staaten. Berliner entomologische Zeitschrift 42 [1897]: 161–288. https://doi.org/10.1002/mmnd.18970420306
- Stein P (1920) Nordamerikanische Anthomyiden. 2.Beitrag. Archiv für Naturgeschichte 84 A 9(1918): е106.
- Tiusanen M, Hebert PDN, Schmidt NM, Roslin T (2016) One fly to rule them all – muscid flies are the key pollinators in the Arctic. Proceedings of the Royal Society, series B 283(1839): е20161271. https://doi.org/10.1098/rspb.2016.1271
- Tizard J, Patel S, Waugh J, Tavares E, Bergmann T, Gill B, Norman J, Christidis L, Scofield P, Haddrath O, Baker A, Lambert D, Millar C (2019) DNA barcoding a unique avifauna: an important tool for evolution, systematics and conservation. BMC Evolutionary Biology 19: e52. https://doi.org/10.1186/s12862-019-1346-y
- Whitworth T, Dawson RD, Magalon H, Beaydry E (2007) DNA barcoding cannot reliably identify species of the blowfly genus Protocalliphora (Diptera: Calliphoridae). Proceedings of the Royal Society – series B 274: 1731–1739. https://doi.org/10.1098/rspb.2007.0062
- Wulp FM van der (1896) Group Anthomyiinae. [Part.] In: Godman FD, Salvin O (Eds) Biologia Centrali-Americana. Zoologia-Insecta-Diptera 2(132), 321–344.
- Xue WQ, Pont AC, Wang MF (2008) A study of the genus Drymeia Meigen (Diptera: Muscidae) from the Tibetan Plateau and surrounding regions of P.R. China, with descriptions of three new species. Proceedings of the Entomological Society of Washington 110: 493–503. https://doi.org/10.4289/07-040.1
- Xue WQ, Wang DD, Wang MF (2009) Three new species of the genus Drymeia Meigen (Diptera: Muscidae) from the Qinghai-Xizang (Tibet) Plateau, China. Entomologica fennica 20: 84–93. https://doi.org/10.33338/ef.84465
- Xue WQ, Wang DD, Zhang XZ, Xiang CQ (2007) Five new species of Drymeia Meigen from the Tibet Plateau, China (Diptera: Muscidae). Zootaxa 1444: 35–51. https://doi.org/10.11646/zootaxa.1444.1.3
Supplementary material
Table S1
Data type: DNA barcoding data set
Explanation note: Details of specimens included in the DNA barcoding data set.