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https://doi.org/10.11646/zootaxa.4848.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:04C94342-9951-4452-9296-AACBD8956113
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by C. Lamas: 6 Jul. 2020; published: 7 Sept. 2020
4848
ZOOTAXA
Magnolia Press
Auckland, New Zealand
Zootaxa 4848 (1): 001–075
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Monograph
The empidoid flies (Diptera: Empidoidea, exclusive of Dolichopodidae) of the
Russian Arctic islands and Svalbard Archipelago
IGOR V. SHAMSHEV1*, BRADLEY J. SINCLAIR2 & OLGA A. KHRULEVA3
1Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg,
199034, Russia
2Canadian National Collection of Insects & Canadian Food Inspection Agency, OPL-Entomology, K.W. Neatby Bldg., C.E.F., 960
Carling Ave., Ottawa, ON, K1A 0C6, Canada
�
bradley.sinclair@canada.ca; https://orcid.org/0000-0001-6413-1606
3A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow, 119071, Russia
�
oa-khruleva@mail.ru; https://orcid.org/0000-0001-6602-0053
*Corresponding author.
�
shamshev@mail.ru; https://orcid.org/0000-0003-3575-4243
SHAMSHEV ET AL.
2 · Zootaxa 4848 (1) © 2020 Magnolia Press
IGOR V. SHAMSHEV, BRADLEY J. SINCLAIR & OLGA A. KHRULEVA
The empidoid flies (Diptera: Empidoidea, exclusive of Dolichopodidae) of the Russian Arctic islands and
Svalbard Archipelago
(Zootaxa 4848)
75 pp.; 30 cm.
7 Sept. 2020
ISBN 978-1-77688-044-7 (paperback)
ISBN 978-1-77688-045-4 (Online edition)
FIRST PUBLISHED IN 2020 BY
Magnolia Press
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 3
Table of Contents
Abstract ...................................................................................................4
Introduction ................................................................................................4
Material and methods ........................................................................................5
Taxonomy .................................................................................................7
Family BRACHYSTOMATIDAE ..............................................................................7
Genus Heleodromia Haliday ..................................................................................7
Heleodromia immaculata Haliday ..............................................................................7
Heleodromia irwini Wagner ...................................................................................7
Heleodromia pullata (Melander) ...............................................................................8
Family EMPIDIDAE ........................................................................................8
Subfamily Clinocerinae ......................................................................................8
Genus Clinocera Meigen .....................................................................................8
Clinocera appendiculata (Zetterstedt) ...........................................................................8
Clinocera nivalis (Zetterstedt) .................................................................................9
Genus Trichoclinocera Collin ................................................................................10
Trichoclinocera lapponica (Ringdahl) ..........................................................................10
Subfamily Empidinae .......................................................................................10
Genus Empis Linnaeus ......................................................................................10
Empis septentrionalis Shamshev & Sinclair sp. nov. ..............................................................10
Genus Rhamphomyia Meigen ................................................................................13
Key to Rhamphomyia of the Russia Arctic islands and Svalbard Archipelago . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Rhamphomyia (Ctenempis) albopilosa Coquillett ................................................................16
Rhamphomyia (Dasyrhamphomyia) brusewitzii Holmgren .........................................................17
Rhamphomyia (Dasyrhamphomyia) erinacioides Malloch .........................................................18
Rhamphomyia (Dasyrhamphomyia) gorodkovi Shamshev, Sinclair & Saigusa sp. nov. ...................................19
Rhamphomyia (Dasyrhamphomyia) hovgaardii Holmgren .........................................................22
Rhamphomyia (Dasyrhamphomyia) mallochi Shamshev, Sinclair & Saigusa sp. nov. ....................................23
Rhamphomyia (Dasyrhamphomyia) nigrita (Zetterstedt) ..........................................................26
Rhamphomyia (Eorhamphomyia) shewelli Sinclair, Vajda, Saigusa & Shamshev . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Rhamphomyia (Lundstroemiella) hybotina (Zetterstedt) ...........................................................28
Rhamphomyia (Pararhamphomyia) armipes Sack ................................................................29
Rhamphomyia (Pararhamphomyia) caudata (Zetterstedt) ..........................................................31
Rhamphomyia (Pararhamphomyia) filicauda Henriksen & Lundbeck ................................................31
Rhamphomyia (Pararhamphomyia) frigida Sinclair, Vajda, Saigusa & Shamshev . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Rhamphomyia (Pararhamphomyia) hoeli Frey ..................................................................32
Rhamphomyia (Pararhamphomyia) kjellmanii Holmgren ..........................................................34
Rhamphomyia (Pararhamphomyia) longestylata Frey ............................................................35
Rhamphomyia (Pararhamphomyia) nordqvistii Holmgren .........................................................37
Rhamphomyia (Pararhamphomyia) septentrionalis Sinclair, Vajda, Saigusa & Shamshev . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Rhamphomyia (Pararhamphomyia) subfilicauda Shamshev & Sinclair sp. nov. ........................................39
Rhamphomyia (Pararhamphomyia) submacrura Shamshev & Sinclair sp. nov. ........................................41
Rhamphomyia (Pararhamphomyia) subtenuiterfilata Shamshev & Sinclair sp. nov. ....................................43
Rhamphomyia (Pararhamphomyia) ursinella Melander ...........................................................45
Rhamphomyia (Pararhamphomyia) wrangeli Shamshev & Sinclair sp. nov. ...........................................46
Rhamphomyia (Rhamphomyia) albosegmentata (Zetterstedt) .......................................................49
Rhamphomyia (Rhamphomyia) morio (Zetterstedt) ...............................................................50
Species unplaced to subgenus .................................................................................51
Rhamphomyia hirtula Zetterstedt .............................................................................51
Rhamphomyia kaninensis Frey ...............................................................................52
Rhamphomyia sublongiseta Shamshev, Sinclair & Saigusa sp. nov. ..................................................54
Rhamphomyia taimyrensis Frey ..............................................................................57
Unnamed species of Rhamphomyia ............................................................................60
Rhamphomyia sp. 1 ........................................................................................60
Family HYBOTIDAE .......................................................................................60
Subfamily Tachydromiinae ...................................................................................60
Genus Platypalpus Macquart .................................................................................60
Platypalpus septentrionalis Shamshev & Sinclair sp. nov. ..........................................................60
Platypalpus subtectifrons Shamshev & Sinclair sp. nov. ...........................................................63
Genus Tachypeza Meigen ....................................................................................66
Tachypeza sericeipalpis Frey .................................................................................66
Discussion ................................................................................................66
Acknowledgements .........................................................................................71
References ................................................................................................72
SHAMSHEV ET AL.
4 · Zootaxa 4848 (1) © 2020 Magnolia Press
Abstract
The empidoid flies of the Russian Arctic islands and Svalbard Archipelago are revised including families Empididae
(4 genera, 32 species), Hybotidae (2 genera, 3 species) and Brachystomatidae (1 genus, 2 species). The following ten
new species are described: Empididae—Empis septentrionalis Shamshev & Sinclair sp. nov. (Russia: Wrangel Island);
Rhamphomyia (Dasyrhamphomyia) gorodkovi Shamshev, Sinclair & Saigusa sp. nov. (Russia: Chukotka mainland, Wrangel
Island, Taymyr, Yakutia; USA: Alaska); R. (D.) mallochi Shamshev, Sinclair & Saigusa sp. nov. (Canada: Yukon; Russia:
Chukotka mainland, Kolyuchin and Wrangel Islands; USA: Alaska); R. (Pararhamphomyia) subfilicauda Shamshev &
Sinclair sp. nov. (Russia: Wrangel Island); R. (P.) submacrura Shamshev & Sinclair sp. nov. (Russia: Wrangel Island);
R. (P.) subtenuiterfilata Shamshev & Sinclair sp. nov. (Russia: Wrangel Island, Yakutia); R. (P.) wrangeli Shamshev
& Sinclair sp. nov. (Russia: Wrangel Island); R. sublongiseta Shamshev, Sinclair & Saigusa sp. nov. (Canada: Yukon;
Russia: Wrangel Island); one species of Rhamphomyia was retained as unnamed due to insufficient material; Hybotidae—
Platypalpus septentrionalis Shamshev & Sinclair sp. nov. (Russia: Wrangel Island), P. subtectifrons Shamshev & Sinclair
sp. nov. (Canada: Northwest Territories, Yukon; Russia: Wrangel Island; USA: Alaska). For previously described species
a short diagnosis and a habitus photo are provided to enable easier recognition. In addition, a key to the species of
Rhamphomyia (Empididae) occurring on the Russia Arctic islands and Svalbard Archipelago is compiled. A lectotype
is designated for Rhamphomyia armipes Sack, 1923. The male and the female of R. taimyrensis Frey and R. kaninensis
Frey are described for the first time, respectively. Tachypeza sericeipalpis Frey is newly recorded from North America
(Canada: Northwest Territories, Nunavut) and first illustration of the male terminalia is provided. Trichoclinocera
lapponica (Ringdahl) is recorded for the first time from the territory of Russia.
Key words: Diptera, Brachystomatidae, Empididae, Hybotidae, taxonomy, new species, Palaearctic, Arctic islands,
tundra zone
Introduction
The superfamily Empidoidea is one of the largest groups of Diptera with about 12 000 described species. The
empidoids are extremely diverse in morphological structures and they occur worldwide (except Antarctica)
inhabiting a very broad range of biotopes (Sinclair & Cumming 2006). The larvae of empidoids breed in a variety
of habitats and are thought to be predators (very exceptionally secondarily phytophagous). Adults are predators or
flower visitors (feeding on nectar and/or pollen), but in some groups predatory activity is associated with the mating
period only.
The classification of higher taxa of Empidoidea proposed by Sinclair & Cumming (2006) is currently commonly
accepted, including herein. It includes five main families, namely Atelestidae, Brachystomatidae, Empididae,
Hybotidae, Dolichopodidae s. lat. (including Microphorinae and Parathalassiinae) and three unassigned genus-
groups (Homalocnemidae, Oreogetonidae and Ragadidae by some authors (e.g., Pape et al. 2011; Wahlberg &
Johanson 2018).
The empidoid flies remain poorly known from the Arctic territories of both Eurasia and North America. They
have been noted in several faunistic and ecological surveys as a marked element of the Arctic biota and are thought
to be important predators and pollinators (Kevan 1972; Danks 1981; Chernov 1995). However, the first revision of
the Arctic fauna of the family Empididae was published only very recently (Sinclair et al. 2019). Although this study
included only the most speciose empidid genus Rhamphomyia Meigen covering the Canadian Arctic Archipelago,
Greenland and Iceland, many species with Holarctic distributions were revealed.
The Eurasian Arctic islands are much less numerous than in North America (primarily the Canadian Arctic
Archipelago), however, they are spread over a vast area extending more than 7000 km from west to east (Fig. 1).
The empidoids of the Eurasian Arctic islands have never been specially studied, with published records rather
scarce. They usually originate from results of sporadic old expeditions and these identifications often need revision.
The first material on the empidoids from the Arctic islands was probably collected by Karl Ernst von Baer, who was
the first naturalist to visit in 1837 the Novaya Zemlya Archipelago (Baer 1838). Some of his specimens are present
in the Collection of the Zoological Institute, St. Petersburg (see below), however, as far as we are aware they have
never been published. Coulson et al. (2014) mentioned the main expeditions that investigated arthropods (including
empidoid flies) from the archipelagos of the Barents Sea. One species of Empididae was recorded from Spitsbergen
(Boheman 1866; Holmgren 1869; Collin 1923). Six species (including four new) were found from Novaya Zemlya
Archipelago and Vaygach Island (Holmgren 1880; Holmgren & Aurivillius 1883; Sack 1923). Nothing has been
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 5
known prior to our study about Empidoidea (exclusive of Dolichopodidae) from the eastern part of the Arctic Ocean;
i.e., from New Siberian Archipelago, Wrangel and other islands. Shamshev (2016) noted several species from these
territories, but without documentary materials that are provided in our paper. Shamshev & Sinclair (2018) revised
recently the type material of Empididae species described by A.E. Holmgren (1880) from Novaya Zemlya.
The purpose of this publication is to summarise primarily taxonomic data on the empidoids of the Russian Arctic
islands and Svalbard Archipelago that is based on a revision of published records and on examination of available
collections (including the type material when it is necessary). Some ecological data were added to supplement
these records. For previously described species a short diagnosis and a habitus photo are provided to enable easier
recognition. In addition, a key to the species of Rhamphomyia (Empididae) occurring on the Russian Arctic islands
and Svalbard Archipelago is compiled.
FIGURE 1. Map of the Russian Arctic islands and Svalbard Archipelago (islands named for only those with records of
empidoids).
Material and methods
This study is based on the Empididae material deposited in the Canadian National Collection of Insects, Ottawa,
ON, Canada (CNC); Czech University of Life Sciences, Prague (CULSP); Finnish Museum of Natural History,
Helsinki (MZH); Naturhistoriska Riksmuseet, Stockholm, Sweden (NHRS); Natural History Museum, University
of Oslo, Oslo, Norway (NHMO); A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of
Sciences, Moscow, Russia; Siberian Zoological Museum, Institute of Systematics and Ecology of Animals of the
Russian Academy of Sciences, Novosibirsk (SZM); Zoological Institute of the Russian Academy of Sciences, St.
Petersburg, Russia (ZIN); Zoological Museum of Moscow University, Moscow (ZMMU).
The material of empidoid flies is rather limited from most islands of the Russian Arctic. The only exception
is Wrangel Island from where quite a large collection was taken by K.B. Gorodkov in 1966, 1971, 1972 (mostly
southern part) and, especially, by O.A. Khruleva during her survey of the arthropods of the entire island (2006–2019).
In addition, L.F. Volkova, U.V. Babiy and A.A. Rodionov (Wrangel Nature Reserve) assisted in O.A. Khruleva’s
field work. O.A. Khruleva collected arthropods from the main biotopes of the island applying different collecting
techniques (i.e., sweep nets, pitfall traps, yellow pan traps). A short description of collecting sites and biotopes (only
where empidoids collected) is given below. The designations of the biotopes (e.g., biotope 1, etc.) refer to these
noted in the material examined sections. The sites are arranged following zonal subdivision of the territory of the
island proposed by Kholod (2013).
1. The northern variant of the arctic tundra subzone. The material was collected by L.F. Volkova in 2016
from two sites on the southeastern part of the island: the vicinity of Thomas Mtn (70°57′N 178°43′Е)—gravelly
SHAMSHEV ET AL.
6 · Zootaxa 4848 (1) © 2020 Magnolia Press
southern slope with spotted herb-dwarf willow-moss cover (biotope 16); the middle flow of Neozhidannaya River
(71°01′N 179°09′Е) (Figs 81, 82), by yellow pan traps—dry high floodplain with forbs-willow-moss cover (biotope
1); gravelly foothill with spotted forbs-willow-dryad (biotope 2) and willow-cereal (biotope 3) cover; wetland with
continuous sedge-moss cover (biotope 4). In 2019, O.A. Khruleva surveyed the following biotopes of the latter site
(yellow pan traps and sweeping): moderately moistened foothill with spotted grass-shrub-moss cover (biotope A)
and with a wormwood-cereal cover (biotope B); wet river terrace with continuous grass-moss cover (biotope W);
low hill with spotted grass-willow-moss cover (biotope G); pebble floodplain with sandy-silt sediments (biotope
D). 2. The southern variant of the Arctic tundra subzone. Some parts of the island located in this variant of the
subzone (environs of Somnitelnaya Bay and Sovetskaya Mtn., upper course of Khischniki River, Rodgers Bay)
were previously studied by K.B. Gorodkov. O.A. Khruleva carried out in 2006 and 2015 a transect from coastal
biotopes on the Southern Plain (70°57′N, 179°38′W) to the slopes of the Somnitelnaya Mtns (70°59′N 179°35′W)
and Mineev Mtns (71°00′N, 179°31′W) (including the valley of the Somnitelnaya River located between them) using
pitfall traps (most part of season) and sweep nets (mid-July). The empidoids were collected only in two biotopes
located in the mountains: on a high floodplain with a willow-herb cover in the upper reaches of the Somnitelnaya
River (biotope 9B) and on the southern slope of the Mineev Mtns with spotted legume-herb-dryad-moss cover
(biotope 14B) (Figs 83, 84). In 2019, the southeastern part of the island was surveyed, with the following sites and
biotopes: the vicinity of Aterton Mtn (70°59′N 178°41′W)—wet base of a hill with a hummocky willow-sedge-moss
cover (biotope B); the vicinity of Rogers Bay (70°59′N 178°29′W)—a foothill, in the dry plot with spotted willow-
moss-herb (biotope A); in a wet depression with spotted willow-sedge-moss cover (biotope 1).
3. The northern variant of the typical tundra subzone. Tundrovaya Mt. (71°18′N, 179°48′W). This site was
surveyed by K.B. Gorodkov in 1972. In 2016, L.F. Volkova collected on the following biotopes (pitfall traps): dry
gravelly base of hill with spotted herb-dryad cover (biotope 2); stream valley with continuous herb-moss cover
(biotope 3); the damp northern slope of hill with wormwood-dwarf willow-moss cover (biotope 5). In 2018, U.V.
Babiy (yellow pan traps): the damp slope of hill with continuous willow-sedge-moss cover (biotope 1); a dry plot
of foothill with sporadic herb-lichen cover (biotope 3); in stream valley with hummocky herb-moss cover (biotope
5), and, in 2019, pebble-loamy floodplain of stream (biotope 2).
The middle flow of Mamontovaya River (71°10′N 179°45′W) (Fig. 85). This site was surveyed by K.B.
Gorodkov in 1966 and 1972. O.A. Khruleva collected the most part of the material in 2006 and 2015 using different
collecting methods. In the river valley flies were collected in dry plots with willow-legume-forb (biotope 2) and
willow-dryad-forb (biotope 1A) cover, as well as in the wetland: osier-bed in the floodplain (biotopes 14, 20A)
and willow-herb-moss cover on the river terrace (biotopes 3, 20). Outside river valley flies were collected in the
zonal habitat—a moderately moisturized low hill with spotted herb-willow-moss cover (biotope 7) and in the more
dry sites with spotted dryad-willow-herb (biotopes 6, 6A, 11) and forb-dryad (biotope 13) cover, as well as in the
tundra-steppe-like plant communities with forbs and xerophilous sedges located on the extremely dry gravelly
slopes of hills and river terraces (biotopes 5, 5A, 10, 17). In addition, flies were collected on Arctic fox burrows with
wormwood-cereal zoogenic meadow. The majority of biotopes were investigated in both seasons. Single specimens
were also collected from this area by A.A. Rodionov in pitfall traps in 2011 on a loamy low hill with hummocky
forb-dryad cover (biotope 4) as well as by O.A. Khruleva in 2019 in the brook Vesely (71°12′N 179°43′W) with
moss-willow-herb cover (biotope A) and in wet base of Inkali Mtn (71°06′N 179°42′W) with hummock dryad-
sedge-moss cover.
The upper flow of Neizvestnaya River. Two sites separated by a distance of nearly 8 km were investigated. The
main upstream (spurs of Pervaya Mtn, 71°09′N, 179°27′W) material was collected by O.A. Khruleva in 2015 (sweep
nets and pitfall traps) in the stream valley with herb-moss-willow cover (biotope 11), on the dry gravel foothill trait
with spotted herb-dryad cover (biotope 12) (Fig. 86), on the dry gravelly southern slope with spotted tundra-steppe
willow-forb-sedge plant communities (biotopes 13, 15) and in the wet runoff slope with dryad-sedge-moss cover
(biotope 14). Single specimens were also collected by O.A. Khruleva in 2019 on a gravelly southern slope with
spotted forb-dryad cover (biotope A). Downstream (71°13′N, 179°19′W) material was collected in 2006 and 2015
using different collecting methods. Empidoid flies were collected on the pebble floodplain (biotope 1A) and in the
osie-bed of the river valley (biotope 4A), different wet habitats with willow-sedge-moss cover (biotopes 5A, 12, 13),
moderately moisturized habitats with dryad-sedge-moss (3A, 3B, 3G), moss-herb-dryad (2, 2B), dryad-cereal-forb
(C 1) and willow-herb (7A) cover. The extreme dry gravelly plots (mainly located on the southern slopes of hills
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 7
and river terrace) occupy tundra-steppe-like communities (biotopes C2, C4, 6, 7B, 10B). The majority of biotopes
were observed in both seasons. Still downstream, in the middle flow of Neizvestnaya River (71°20′N 179°30′W)
flies were collected by U.V. Babiy in yellow pan traps in 2018 and 2019 on the pebble floodplain (biotope 1) and in
the osier-bed in the river valley (biotope 2).
The specimens collected by yellow pan traps and pitfall traps were kept in alcohol and later dried and pinned.
Digital images were taken with a Nikon SMZ 1500 stereomicroscope equipped with Nikon D700 digital SLR
camera. Terms used for adult structures follow those of Cumming & Wood (2017).
The following abbreviations are used in the descriptions and material examined sections: acr—acrostichal
setae, dc—dorsocentral setae, npl—notopleural setae, pal—postalar setae, pprn—postpronotal setae, presut spal—
presutural supra-alar setae, psut spal—postsutural supra-alar setae; BT—biotope, PT—pitfall traps, Sw—sweeping,
YPT—yellow pan traps; most frequent collectors: KBG—K.B. Gorodkov, OAK—O.A. Khruleva, LFV—L.F.
Volkova: administrative unites of the Russian Federation: AO—Autonomous Okrug, Prov.—Province (= oblast),
Terr.—Territory (= kray).
Label data for primary types are cited from the top downward, with the data from each label in quotation
marks. Labels are cited in full, with original spelling, punctuation, and date, and label lines are delimited by a slash
(/). Additional information is included in square [ ] brackets. The repository of each type is given in parentheses.
Secondary type data are abridged and listed alphabetically.
The taxa are arranged hierarchically and alphabetically according to the categories of family, subfamily,
genus, subgenus, and species. The material originating from the same island is arranged by locality and is listed
chronologically (on Wrangel Island according to the zonal subdivision noted above). The species of Rhamphomyia
are organized alphabetically by subgenus following Sinclair et al. (2019). Transliteration of Cyrillic (Russian) names
follows those of Shamshev (2016). The survey in the Russian Arctic is restricted to any marine (oceanic) island
independent of offshore distance or latitude. Distribution range beyond the Russian Arctic islands and Svalbard
Archipelago is provided, based on Shamshev (2016) and Sinclair et al. (2019).
Taxonomy
Family BRACHYSTOMATIDAE
Genus Heleodromia Haliday
Heleodromia immaculata Haliday
Heleodromia immaculata Haliday, 1833: 159. Type locality: Ireland, Holywood, Downshire.
Sciodromia immaculata: Sack, 1923: 7 (expedition results).
Remarks. Heleodromia immaculata was reported from Novaya Zemlya by Sack (1923), but this record needs to be
confirmed. Species identification of Heleodromia is dependent on careful study of the complex male terminalia and
the shape of the male tergite 8 (Sinclair et al. 2011).
Heleodromia irwini Wagner
Heleodromia irwini Wagner, 1985: 37. Type locality: Scotland, Moray, Dorback Burn, UK.
Other references: Sinclair et al., 2011: 641 (revision, redescription); Shamshev, 2016: 17 (checklist).
Material examined. RUSSIA. Chukotka AO (Wrangel Island): middle flow of river Neozhidannaya, 71°01′N
179°09′E, YPT, BT 4, 12–16.vii.2016, LFV (1 ♂, ZIN).
Recognition. Small greyish delicate flies (body length 3–3.5 mm) with very slender legs almost lacking distinct
setae, eyes dichoptic in both sexes, separated by broad V-shaped flattened frons; legs black to reddish brown,
“knees” often somewhat paler; habitually very similar to H. pullata (Fig. 2, see below). Male terminalia large,
reniform; tergite 8 not broader than sternite 8, only produced medially as a shallow bilobed to forked projection
SHAMSHEV ET AL.
8 · Zootaxa 4848 (1) © 2020 Magnolia Press
and lacking posterolateral projection (Sinclair et al. 2011: 641, fig. 10); epandrial lobe slender, finger-like, bearing
apical seta; surstylus slender, finger-like, with scattered setae, with small preapical thumb-like projection medially;
apex with short dentiform process; phallus with slender, needle-like apex.
Distribution. Holarctic. The distribution of this species appears to be represented by widely disjunct populations
exhibiting boreo-alpine connections in Europe and North America (Plant 2005; Plant et al. 2017; Sinclair et al.
2011). Our record of H. irwini from Wrangel Island may support this suggestion. In Eurasia, this species is known
from Austria, Germany, Italy, Switzerland, Scotland and the Far East of Russia (Amurskaya and Magadanskaya
Provinces); in North America, it is recorded from western part of Canada and USA. It seems that H. irwini is found
here for the first time from the tundra zone.
Habitat. The species was collected in wetlands in a river valley.
Heleodromia pullata (Melander)
(Fig. 2)
Sciodromia pullata Melander, 1902: 345. Type locality (by lectotype designation): Las Vegas Range, New Mexico, USA.
Other references: Sinclair et al., 2011: 642 (lectotype designation, redescription); Shamshev, 2016: 17 (checklist).
Material examined. RUSSIA. Chukotka AO (Wrangel Island): Somnitelnaya Bay, on stony river shore,
23.vii.1971, KBG (1 ♂, ZIN); upper flow of river Khischniki, 7 km SE Sovetskaya Mtn, 200 m, dryad tundra
near river, 11.vii.1972, KBG (3 ♂, 1 ♀, ZIN); middle flow of river Mamontovaya, 71°10′N 179°45′W, BT 2, YPT,
2–4.vii.2015, OAK (2 ♂, 2 ♀, ZIN); same locality, 71°10′N 179°45′W, BT 3, Sw, 1.vii.2015, OAK (1 ♀, ZIN);
upper flow of river Neizvestnaya, 71°13′N 179°19′W, BT 2В, 8–29.vii.2015, OAK (1 ♂, 1 ♀, ZIN); same locality,
71°13′N 179°19′W, BT 13, YPT, 6–15.vii.2015, OAK (1 ♂, ZIN).
Recognition. Small greyish delicate flies (body length 3–3.5 mm) with very slender legs almost lacking distinct
setae, eyes dichoptic in both sexes, separated by broad V-shaped flattened frons; legs black to reddish brown, “knees”
often somewhat paler (Fig. 2). Male terminalia large, reniform; tergite 8 simple, lacking dorsomedial elongation
or process (Sinclair et al. 2011: 644, fig. 11B); epandrial lobe slender, digitiform; surstylus slightly shorter than
epandrial lobe, with apical third slender and digitiform; phallus with apex membranous and broad.
Distribution. Holarctic; in North America, H. pullata is widely distributed in northern Canada and Alaska,
occurring on tundra and in boreal forests, with isolated populations in the Rocky Mountains of Arizona, Colorado
and New Mexico (Sinclair et al. 2011); in Eurasia, it is known from Taymyr Peninsula and Magadanskaya Province
of Russia. Among the Russian Arctic islands, H. pullata is recorded only from Wrangel Island.
Habitat. This species was collected on Wrangel Island mainly in river beds, where it was found in biotopes with
different moisture levels (also on stones).
Family EMPIDIDAE
Subfamily Clinocerinae
Genus Clinocera Meigen
Clinocera appendiculata (Zetterstedt)
Wiedemannia appendiculata Zetterstedt, 1838: 559. Type locality: Schiervöe, Nordland, Norway.
Other references: Sack, 1923: 7 (expedition results); Sinclair, 1999: 224 (revision); Shamshev, 2016: 18 (checklist).
Remarks. The identification history of C. appendiculata is somewhat confusing (see Sinclair (1999) for details),
consequently the record of this species from Novaya Zemlya (Sack 1923) requires confirmation. We expect this
record to possibly be Clinocera aucta (Zetterstedt, 1849) (Sinclair 2008).
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 9
FIGURES 2–5. Brachystomatidae (2) and Empididae, Clinocerinae (3–5). 2. Heleodromia pullata (Melander), male habitus,
lateral view; 3. Clinocera nivalis (Zetterstedt), male habitus, lateral view; 4. Trichoclinocera lapponica (Ringdahl), male habitus,
lateral view; 5. same species, male postabdomen, lateral view.
Clinocera nivalis (Zetterstedt)
(Fig. 3)
Hemerodromia nivalis Zetterstedt, 1838: 545. Type locality: Lapland, (?)Sweden.
Other references: Sinclair, 2008: 174 (revision, redescription); Shamshev, 2016: 19 (checklist).
SHAMSHEV ET AL.
10 · Zootaxa 4848 (1) © 2020 Magnolia Press
Material examined. RUSSIA. Archangelskaya Prov. (Novaya Zemlya Archipelago): Southern coast near Cape
Cherny, 21, 29.vii.1911, Rusakov (1 ♂, 2 ♀, ZIN); Eastern coast, Karskie vorota—Matochkin Shar, viii–ix, Rusakov
(1 ♂, 1 ♀, ZIN); Matochkin Shar, brook Nochuev, 22.v.1925 (1 ♀, ZIN), Vakulenko; Matochkin Shar, 10.vii.1925,
Vakulenko (1 ♀, ZIN). Chukotka AO (Wrangel Island): SW mountain Sovetskaya, upper flow of Khrustalnaya
River, under snowfield on detritus, 3.viii.1971, KBG (1 ♀, ZIN); Mineev Mtns, 5 km N Somnitelnaya, slope, swamp
spring, 26.vii.1971, KBG (1 ♂, 1 ♀, ZIN); lower flow of Khischniki River, foothills of Mineev Mtns, swamp tundra,
11.vii.1972, KBG (1 ♂, ZIN); Rogers Bay, small swamp, on surface of water, 20.vii.1971, KBG (2 ♂, 1 ♀, ZIN).
Recognition. This species of Clinocera is readily distinguished by the presence of an auxiliary crossvein
between R2+3 and R4, lower half of face with silvery pruinescence and a dark ring around the anterior spiracle (Fig.
3). Distribution. A Holarctic species distributed across the subarctic and arctic areas of Eurasia and North America
(Sinclair 2008). In Russia, C. nivalis is known from Murmanskaya Province, Novaya Zemlya, north of Yakutia,
Chukotka and Wrangel Island.
Habitat. On Wrangel Island, this species was collected in various wet habitats.
Genus Trichoclinocera Collin
Trichoclinocera lapponica (Ringdahl)
(Figs 4, 5)
Atalanta lapponica Ringdahl, 1933: 281. Type locality (by lectotype designation): Tjuonjatjakko, Sweden.
Other references: Chvála & Wagner, 1989: 335 (catalogue).
Trichoclinocera lapponica: Sinclair, 1994: 1023 (revision, lectotype designation, redescription).
Material examined. CANADA. Quebec [new records since Sinclair (1994)]: 10 km SW Tasiujaq, trib. of Finger
River, 17.viii.1996, B.J. Sinclair (7 ♂, CNC). RUSSIA. Chukotka AO (Wrangel Island): middle flow of
Neozhidannaya River, 71°01′N, 179°09′W, BT, YPT, 8–9.vii.2019, OAK (1 ♀, ZIN); Somnitelnaya River, on water,
water temperature +5, air temperature +2.5, 3.viii.1971, KBG (10 ♂, 3 ♀, ZIN); same locality, under pebble near
water, 3.viii.1971, KBG (2 ♂, 1 ♀, ZIN); SW Sovetskaya Mtn, upper flow of Khrustalnaya River, 3.viii.1971,
KBG (2 ♂, 12 ♀, ZIN); same locality, snowfield, on water, air temperature +3, water temperature +7, snowfall,
4.viii.1971, KBG, Chelnokov (1 ♂, 7 ♀, ZIN); environs of Tundrovaya Mtn, 71°17.863′N, 179°47.875′W, BT 2,
YPT, 7–11.vii.2019, U.V. Babiy (1 ♀, ZIN); middle flow of Mamontovaya River, N Perkatkun, on emergent stone
in river, 29.vii.1972, KBG (3 ♂, 1 ♀, ZIN).
Recognition. Among clinocerines, Trichoclinocera is distinguished by the presence of setae on vein R1. This
species is distinguished readily by its dorsoventrally flattened tarsomeres 4 and 5, convex face and small eyes (Figs
4, 5).
Distribution. Holarctic species known from northern Finland and Sweden as well as it is widespread throughout
the arctic region of North America (Sinclair 1994); recorded for the first time from the territory of Russia.
Habitat. On Wrangel Island, this species was collected mainly on pebble floodplains, near water.
Subfamily Empidinae
Genus Empis Linnaeus
Empis septentrionalis Shamshev & Sinclair sp. nov.
(Figs 6, 7)
urn:lsid:zoobank.org:act:1E05618C-B300-4E56-B381-1A0865B7462E
Type material. HOLOTYPE ♂, labelled: [RUSSIA. Chukotka AO (Wrangel Island)] “[printed in Cyrillic,
Russian] Chukotskiy AO [= Autonomous Okrug] o. [= ostrov, island] Wrangel/ Verch. tech. [= verkhnee techenie,
upper flow] river Neizvest-/ naya 71°13′N 179°19′W/ O.A. Khruleva 8.7.2015/ 2B”; “Holotypus/ Empis/
septentrionalis/ Shamshev, Sinclair sp. nov. [red label]” (INS_DIP_0000612, ZIN).
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 11
PARATYPES: RUSSIA. Chukotka AO (Wrangel Island): same locality as holotype, BT 7A, 5.vii.2006,
OAK (1 ♂, 1 ♀, CNC); same locality as holotype, BT 7A, 10.vii.2015, OAK (1 ♂, ZIN); same locality as holotype,
BT 7B, 10.vii.2015, OAK (7 ♀, ZIN); same locality as holotype, BT C1, YPT, 4–15.vii.2015, OAK (1 ♂, ZIN);
same locality as holotype, BT 1A, 6–13.vii.2015, OAK (1 ♂, ZIN); Mineev Mtns, 71°00′N 179°31′W, BT 14B,
14.vii.2006, OAK (1 ♂, ZIN); environs of Pervaya Mtn (71°10′N 179°27′W), BT A, hand collecting, 21.vii.2019,
OAK (1 ♀, ZIN).
Diagnosis. Mid-sized (wing length 5–5.5 mm) robust flies with brownish pruinescent thorax and abdomen
and almost entirely dark brown legs. Male dichoptic; occiput slightly convex with numerous long, flattened setae
laterally behind eye; thorax black setose, scutum with 4 vittae; legs with only hind tarsomeres 1–3 brownish yellow,
hind femur somewhat thickened, hind tibia very slender; apical part of phallus uniformly tubular. Female with
brownish infuscate wing; abdomen densely brown pruinescent on tergites dorsally.
Description. Body length 5.2–5.9 (holotype 5.3); wing length 5–5.5 (holotype 5.1) mm. Male (Fig. 6). Head
with dense brownish pruinescence on face, frons, postgena, ocellar triangle and occiput; entirely dark setose; clypeus
shiny. Dichoptic; ommatidia equally small. Frons broad, below ocellar tubercle broader than distance between outer
margins of posterior ocelli; with marginal setulae. Face broad, bare. Ocellar triangle with several long fine setae.
Occiput slightly convex laterally behind eyes; with numerous long mostly flattened setae on convex part, fine long
postoculars and similar setae on upper part; postgena with numerous hair-like setae. Antenna black; scape short,
slightly longer than pedicel, both with short setulae; postpedicel conical, with slightly concave ventral margin on
apical part, nearly 2X longer than wide; stylus rather long, slightly longer (1.3–1.4X) than postpedicel basal width.
Palpus dark (sometimes yellowish, 2 ♂); with several black moderately long fine setae. Proboscis with labrum
mostly brownish, reddish-brown apically, about 1.5X longer than head height.
Thorax dark in ground-colour, brownish pruinescent; with black setation; scutum with 4 indistinct narrow
blackish brown vittae (dorsal view). Proepisternum with tuft of numerous mostly long and slightly flattened setae
(with some additional shorter fine setae) on lower section; upper proepisternum in front of spiracle bare. Prosternum
bare. Antepronotum with numerous short fine setae. Postpronotal lobe covered with very numerous subequally
long setae. Mesonotum with hardly differentiated setae; several long fine intra-alars just behind postpronotal lobe;
supra-alar space with several similar setae before suture and less numerous setae behind suture, notopleuron with
numerous long setae, some posterior setae somewhat stronger; 2 long strong pal (sometimes with additional setulae),
8–10 sctl; acr long, fine, arranged in 2 irregular rows, lacking on prescutellar depression; presutural dc similar to acr,
multiserial, postsutural dc less numerous, 3–4 prescutellars longest. Laterotergite with numerous long fine setae.
Anterior and posterior spiracles brown.
Legs robust, almost entirely dark brown, only hind tarsomeres 1–3 (sometimes 1–4) brownish yellow; black
setose; hind femur somewhat thickened, mid tibia slightly arcuate, hind tibia very slender. Coxae and trochanters
with simple setae. Femora whitish pilose ventrally (less distinct on fore femur); fore femur with long fine setae
anteroventrally, posteroventrally and posteriorly; mid femur with numerous spine-like setae of different lengths
ventrally, posteroventrals longer, some moderately long anterodorsal setae; hind femur with numerous long flattened
setae anterodorsally (except short subapical part), only pale pilose along lower half anteriorly (except several setae
near extreme base), rows of long, strong anteroventral and posteroventral setae, short spine-like setae ventrally.
Tibiae devoid of strong setae (including setae of subapical circlet); fore tibia with some short, fine setae posteriorly;
mid tibia with erect setulae ventrally; hind tibia pale pilose ventrally, no seta in posteroapical comb. Hind tarsomeres
with somewhat finer and sparser setulae.
Wing membrane very faintly infuscate, with brownish veins; CuA+CuP (anal vein) complete; cell dm with
elongate apex. Pterostigma brownish yellow. Basal costal seta absent. Anal lobe well-developed; axillary incision
acute. Squama dirty yellow, brownish fringed. Halter with yellow knob and brownish stem.
Abdomen dark, brownish pruinescent (somewhat denser on tergites laterally and sternites); covered with
numerous long, dark fine setae (shorter on tergites dorsally). Segment 8 with tergite and sternite separated; tergite 8
mostly membranous, represented by two weakly sclerotized, narrow lateral sclerites each bearing 0–1 setae; sternite
simple, nearly as long as sternite 7 (lateral view), with numerous long setae along posterior margin.
Terminalia (Fig. 7). Cerci broadly yellowish to brownish yellow, narrowly brownish along upper margin and
anteriorly; separated from each other and from epandrium; cercus elongate oval (lateral view), with rounded apex,
slightly extended beyond apex of epandrium, covered with short dark setae. Epandrium almost entirely brownish,
narrowly yellowish brown along lower margin; subtriangular (lateral view), with dark setae more numerous and
longer along lower margin. Hypandrium mostly membranous, rim-like sclerotized along upper margin, bare. Phallus
SHAMSHEV ET AL.
12 · Zootaxa 4848 (1) © 2020 Magnolia Press
almost uniformly broad, slightly narrowed near middle, S-like bend, sulcate on subapical portion, with pair of small
projections near apex beneath. Ejaculatory apodeme extended far beyond basal curvature of phallus, with subequal
lateral and vertical wings and somewhat narrower lower wing.
Female. Occiput not convex laterally, without flattened setae, ocellar tubercle with short setae; palpus somewhat
paler, yellowish brown, sometimes with yellowish apex or yellowish. Thorax and legs with shorter setation; hind
femur not thickened, hind tibia simple; hind femur without flattened setae and pale pilosity along lower half
anteriorly, entirely covered with setulae anteriorly; fore tibia without fine setae posteriorly, mid and hind tibiae with
usual setulae ventrally; hind tarsus brownish, with similar setation to fore and mid tarsi. Wing darker, brownish
infuscate, pterostigma brownish. Abdomen densely brownish grey pruinescent on tergites laterally and sternites,
brown pruinescent on tergites dorsally (except noted); tergites 7–8 extensively shiny (except posterior margin),
also sternites 7–8 usually subshiny to shiny anteriorly; only tergites 1–2 with some short setae laterally, otherwise
tergites covered with scattered setulae; cercus black, with dark setulae.
Distribution. Palaearctic, known only from Wrangel Island.
Etymology. The specific name is from the Latin septentrionalis (north, northern), in reference to the distribution
of this species.
FIGURES 6–8. Empis. 6. E. septentrionalis sp. nov., holotype male, habitus, lateral view; 7. same species, male terminalia,
lateral view; 8. E. jacutiensis Shamshev, male terminalia, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory
apodeme; epand—epandrium; hypd—hypandrium; ph—phallus; tg 8—tergite 8.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 13
Remarks. Empis septentrionalis sp. nov. belongs to the group of species with uncertain subgeneric position
within Empis (Shamshev 2001a). The group is known almost exclusively from the Asiatic part of the Palaearctic
(except one species) and is especially diverse in Middle Asia. In addition, three undescribed species are known from
Yukon in Canada. The new species is very similar to E. jacutiensis Shamshev occurring in Yakutia and Chukotka
of Russia (Shamshev 2016). Empis septentrionalis sp. nov. can be readily distinguished from E. jacutiensis by
uniformly tubular apical part of the phallus (cup-shaped apically in E. jacutiensis) (Fig. 8) and dark brown hind
tibia (yellowish to brownish yellow in E. jacutiensis). In addition, the male of E. septentrionalis sp. nov. usually has
dark palpi (yellowish in E. jacutiensis), but this character appears to be somewhat variable. The female of the new
species differs from the female of E. jacutiensis by dark brown hind tibia and tarsus and by brown pruinescence on
abdominal tergites dorsally (abdomen uniformly light grey pruinescent in the female of E. jacutiensis).
Habitat. Almost all specimens were collected in the warmer central region of Wrangel Island, mainly in dry
habitats.
Genus Rhamphomyia Meigen
Key to Rhamphomyia of the Russia Arctic islands and Svalbard Archipelago
The key provided below follows generally the key compiled by Sinclair et al. (2019) to species of the Canadian
Arctic Archipelago, Greenland and Iceland and we recommend that this publication be consulted for additional
species that may be found in the Eurasian Arctic islands. Identification of Asiatic species of Rhamphomyia remains
problematic due to very high diversity and weak species group definitions and classification. Barták & Kubík (2009)
and Saigusa (2012) should be consulted for keys to species groups of Rhamphomyia and references to available
revisions.
1 Male (unknown in R. nordqvistii, R. sp. 1) ..................................................................2
- Female (unknown in R. armipes, R. submacrura sp. nov., R. subfilicauda sp. nov.) ................................28
2 Axillary lobe of wing very little developed, axillary excision extremely obtuse (Fig. 27); eyes dichoptic, frons shiny [Additional
characters: occiput shiny on upper part, greyish pruinescent on middle.] . . . . . . .R. (Lundstroemiella) hybotina (Zetterstedt)
- Axillary lobe of wing well developed, axillary excision at most 90°; eyes holoptic or frons pruinescent . . . . . . . . . . . . . . . . . 3
3 Prosternum clothed with long white hair-like setae (Sinclair et al. 2019: fig. 3D); sternite 6 with cluster of strong golden setae;
sternite 7 with pair of horn-like projections ventrally (Fig. 9) . . . . . . . . . . . . . . . . . . . . . .R. (Ctenempis) albopilosa Coquillett
- Prosternum bare; sternite 6 without cluster of golden setae; sternite 7 unmodified or modified different than above . . . . . . . . 4
4 Cerci strongly prolonged anteriorly, extended at least to tergite 6 (Figs 10–13) (subgenus Dasyrhamphomyia) ............5
- Cerci not extended anteriorly beyond tergite 8 .............................................................10
5 Halter yellow ........................................................................................6
- Halter brown .........................................................................................8
6 Cerci very long, extended nearly to middle of abdominal tergite 3 (Fig. 13); hind femur without setae on about apical 2/3
ventrally, only densely pale pilose ....................................................R. (D.) nigrita Zetterstedt
- Cerci much shorter, extended at most to abdominal tergite 5; hind femur with numerous short setae over entire length
ventrally ............................................................................................7
7 Hind tibia thickened; wing whitish (Fig. 14); epandrium brownish to reddish brown (Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . .
.....................................................R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.
- Hind tibia slender; wing faintly infuscate (Fig. 20); epandrium yellowish (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
......................................................R. (D.) mallochi Shamshev, Sinclair & Saigusa sp. nov.
8 Frons very broad, in middle nearly as broad as distance between outer margins of posterior ocelli, almost parallel-sided; hind
femur without setae on about apical 3/4 ventrally, only densely pale pilose (Fig. 11) . . . . . . . . . .R. (D.) erinacioides Malloch
- Frons very narrow on middle part, at most slightly broader than anterior ocellus or eyes touching; hind femur with numerous
short setae over entire length ventrally .....................................................................9
9 Mesonotal setae strong, bristle-like (Fig. 10); eyes separated by very narrow frons . . . . . . . . . . . R. (D.) brusewitzii Holmgren
- Mesonotal setae almost uniformly fine, hair-like (Fig. 12); eyes usually touching . . . . . . . . . . . . R. (D.) hovgaardii Holmgren
10 Subepandrial sclerite prolonged into pair of medioposterior lobes beneath cerci (Fig. 31) (Sinclair et al. 2019: figs 25A, C) 11
- Cercus without pair of subepandrial lobes beneath ..........................................................14
11 Hind femur with triangular posteroventral projection beyond mid-length; hind tibia with triangular posteroventral projection on
basal third (Fig. 30) .....................................................R. (Pararhamphomyia) armipes Sack
- Hind femur and tibia without projections ..................................................................12
12 Abdomen covered with black setae; fore coxa with thin setae anteriorly; hind tibia with 4–5 anterodorsal and several posterodorsal
prominent setae, hind basitarsus with only short setae dorsally (Sinclair et al. 2019: fig. 34B); wing membrane whitish (Fig.
34) ....................................................................................R. (P.) hoeli Frey
SHAMSHEV ET AL.
14 · Zootaxa 4848 (1) © 2020 Magnolia Press
- Abdomen covered with pale setae; fore coxa with strong blunt-tipped setae anteriorly; hind tibia and basitarsus covered with
long, dense, sometimes woolly, setae dorsally; wing membrane hyaline to slightly infuscate . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Phallus forming broad loops (Sinclair et al. 2019: fig. 36C); hind tibia clavate (Fig. 35) . . . . . . . .R. (P.) kjellmanii Holmgren
- Phallus not forming loops, only gently curved (Sinclair et al. 2019: fig. 25A, C); hind tibia more slender, evenly thickened
towards apex (Fig. 33) ...........................................................R. (P.) frigida Sinclair et al.
14 Abdominal segments 7 and 8 asymmetrical, strongly sclerotized (somewhat darker) than preceded segments, segment 7
enlarged; terminalia slightly rotated to right (Fig. 52). [Additional characters: hind tibia strongly curved inward near basal third
(posterior view) (Fig. 49); phallus mostly hidden within epandrium, hook-like curved to left at apex (Fig. 53)] . . . . . . . . . . . .
..............................................................R. (P.) wrangeli Shamshev & Sinclair sp. nov.
- Abdominal segments symmetrical, normally sclerotized, segment 7 unmodified; terminalia unrotated . . . . . . . . . . . . . . . . . 15
15 Upper proepisternum in front of spiracle with setae; basal costal seta usually present (often 2 setae in R. shewelli or sometimes
indistinct in R. sublongiseta sp. nov.) ....................................................................16
- Upper proepisternum in front of spiracle without setae; basal costal seta usually absent (except R. taimyrensis) ..........21
16 Wing axillary incision very acute, at most 45°. Cercus without notch dorsally, at most tuberculate; phallus short, gently
curved .............................................................................................17
- Wing axillary incision more than 45°, usually close to 90°. Cercus with notch dorsally (Fig. 60); phallus long, sinuate . . . . 19
17 Halter yellow; abdomen pruinose dorsally and narrowly shiny laterally (Fig. 26) R. (Eorhamphomyia) shewelli Sinclair et al.
- Halter brown; abdomen entirely pruinose, although sometimes thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Face with several fine setae; abdominal tergites faintly pruinescent, somewhat lustrous (Fig. 53) . . . . . . . . . . . . . . . . . . . . . . .
....................................................................R. (Rhamphomyia) morio (Zetterstedt)
- Face bare; abdominal tergites densely light grey pruinescent (Fig. 52) . . . . . . . . . . . . . . . R. (R.) albosegmentata (Zetterstedt)
19 Hind basitarsus strongly expanded (Fig. 62); cercus with notch forming small projection slightly beyond middle dorsally (Fig.
64) .....................................................R. sublongiseta Shamshev, Sinclair & Saigusa sp. nov.
- Hind basitarsus slender; cercus with finger-like projection near base dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Wing somewhat whitish (Fig. 57); mid tibia and basitarsus with numerous very long setae dorsally (2–2.5X longer than tibia
width) (Fig. 58) ........................................................................R. kaninensis Frey
- Wing faintly infuscate; mid tibia and basitarsus with short setae (at most as long as tibia width) (Fig. 56) . . . . . . . . . . . . . . . . .
..................................................................................R. hirtula Zetterstedt
21 Phallus with small, narrow U-shaped loop on subapical part (Fig. 67); cercus about as long as epandrium, with straight dorsal
margin. [Additional character: hind tibia curved inward closer to base (posterior view)] (Fig. 66) . . . . . . . R. taimyrensis Frey
- Phallus gradually curved; cercus distinctly shorter than epandrium, usually more or less concave dorsally . . . . . . . . . . . . . . 22
22 Anal vein reaching wing margin, entirely sclerotized (Fig. 43). [Additional characters: halter brownish; mesonotum and
abdomen with only black setae; scutum entirely pruinescent; hind tibia straight; hind basitarsus slightly expanded, cylindrical,
covered with short setae; epandrium nearly 4X longer than cercus (Figs 43, 44).] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
...........................................................R. (P.) submacrura Shamshev & Sinclair sp. nov.
- Anal vein disappearing before wing margin, at most evanescent (or fold-like) apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 Halter with yellow knob; mesonotum (except stronger setae) and abdomen covered with pale thin setae; hind tibia curved
(posterior view); hind basitarsus spindle-shaped, convex dorsally, longer than half length of tibia (Fig. 39). [Additional
characters: scutum with 4 shiny vittae; hind basitarsus covered with numerous long setae dorsally; epandrium nearly 1.5X
longer than cercus.] ...................................................... R. (P.) septentrionalis Sinclair et al.
- Halter brown; mesonotum and abdomen with black setation; hind tibia straight; hind basitarsus cylindrical or clavate, shorter
than half length of tibia ................................................................................24
24 Anepisternum shiny ..................................................................................25
- Anepisternum pruinose ................................................................................26
25 Scutum faintly greyish pruinescent, with two shiny median vittae (sometimes additional less distinct vitta present on supra-alar
space); hind tibia with uniformly long setae dorsally; hind basitarsus clavate (Fig. 48); cercus spatulate beyond median cavity
(lateral view) ....................................................................R. (P.) ursinella Melander
- Scutum shiny, at most very faintly pruinescent in front of scutellum; hind tibia with long setae only near apical half dorsally;
hind basitarsus more or less cylindrical; cercus finger-like beyond median cavity (lateral view) (Figs 36, 37) . . . . . . . . . . . . . .
................................................................................R. (P.) longestylata Frey
26 Acrostichals and presutural dorsocentrals very short, fine, the latter 1–2-serial; mesonotum and abdomen almost shiny.
[Additional characters: fore and mid tibiae and tarsi with short setae, hind tibia with moderately long setae dorsally; hind
basitarsus clavate, tarsomere 2 thickened, both with long, dense fine setae dorsally (Fig. 45).] . . . . . . . . . . . . . . . . . . . . . . . . . .
....................................................... R. (P.) subtenuiterfilata Shamshev & Sinclair sp. nov.
- Acrostichals and presutural dorsocentrals long, strong, the latter at least 3–4-serial; mesonotum and abdomen densely
pruinose ...........................................................................................27
27 Smaller, wing 4.3 mm; hind tibia with long dense fine setae dorsally; wing somewhat whitish (Fig. 41) . . . . . . . . . . . . . . . . . .
...........................................................R. (P.) subfilicauda Shamshev & Sinclair sp. nov.
- Larger, wing 4.8–5.4 mm; hind tibia with bristle-like setae dorsally; wing faintly infuscate (Fig. 32) . . . . . . . . . . . . . . . . . . . . .
..................................................................R. (P.) filicauda Henriksen & Lundbeck
28 Axillary lobe of wing almost absent, axillary excision extremely obtuse; frons shiny. [Additional characters: occiput shiny on
upper part, greyish pruinescent on middle.] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .R. (L.) hybotina (Zetterstedt)
- Axillary lobe of wing well-developed, axillary excision at most slightly more than 90°; frons pruinescent . . . . . . . . . . . . . . 29
29 Prosternum clothed with long white hair-like setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .R. (C.) albopilosa Coquillett
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 15
- Prosternum bare .....................................................................................30
30 Anal vein (CuA+CuP) reaching wing margin, usually entirely sclerotized (sometimes slightly weaker about middle) (variable
in R. erinacioides) ...................................................................................31
- Anal vein (CuA+CuP) not reaching wing margin or evanescent (fold-like) apically (variable in R. hoeli) ...............44
31 Basal costal seta present (in R. shewelli often 2 setae) ........................................................32
- Basal costal seta absent ...............................................................................38
32 Upper proepisternum in front of spiracle without setae; halter with yellow knob; mid and hind femora with anterodorsal and
posteroventral rows of pennate setae (except extreme base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. taimyrensis Frey
- Upper proepisternum in front of spiracle with setae; halter brown; legs with simple setae . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 Abdomen pruinose dorsally and narrowly shiny laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. (E.) shewelli Sinclair et al.
- Abdominal tergites uniformly pruinose ...................................................................34
34 Face with several fine setae ........................................................R. (R.) morio (Zetterstedt)
- Face bare ...........................................................................................35
35 Acr and dc strong bristle-like; scape and pedicel slightly paler than postpedicel; thorax lighter, grey pruinose . . . . . . . . . . . . .
......................................................................R. (R.) albosegmentata (Zetterstedt)
- Acr and dc fine; antenna uniformly dark; thorax darker, brownish pruinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 Anal lobe of wing darker than wing tip; hind tibia and basitarsus slightly thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.......................................................R. sublongiseta Shamshev, Sinclair & Saigusa sp. nov.
- Wing membrane uniformly infuscate; at least hind basitarsus slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
37 Hind tibia slender, entirely short setose; abdomen lighter, densely greyish pruinose . . . . . . . . . . . . . . . . . . R. kaninensis Frey
- Hind tibia somewhat dilated, almost bare anteriorly; abdomen darker, densely brownish pruinose . . . . . R. hirtula Zetterstedt
38 Legs with pennate setae, at least mid and hind femora (somewhat variable in R. gorodkovi sp. nov.) ...................39
- Legs covered with simple setae, at most sometimes some setae on mid and hind femora slightly flattened . . . . . . . . . . . . . . 41
39 Halter brown; hind tibia with pennate setae dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. (D.) hovgaardii Holmgren
- Halter with yellow knob; hind tibia with simple setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40 Hind tibia evenly broadened toward apex [see also couplet 42] . . . R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.
- Hind tibia slender ........................................R. (D.) mallochi Shamshev, Sinclair & Saigusa sp. nov.
41 Halter at least with yellow knob .........................................................................42
- Halter brown ........................................................................................43
42 Legs uniformly dark, hind tibia slender; wing membrane brownish . . . . . . . . . . . . . . . . . . . . . . . . . .R. (D.) nigrita Zetterstedt
- Legs yellowish to brownish yellow, hind tibia evenly broadened toward apex; wing membrane faintly infuscate . . . . . . . . . . .
.....................................................R. (D.) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.
43 Smaller, wing length about 5 mm; mesonotum dark brown (dorsal view); presutural dorsocentrals multiserial . . . . . . . . . . . . .
.............................................................................R. (D.) erinacioides Malloch
- Larger, wing length 6–6.5 mm; mesonotum greyish (dorsal view); presutural dorsocentrals biserial . . . . . . . . . . . . . . . . . . . . .
............................................................................R. (D.) brusewitzii Holmgren
44 Abdomen covered with pale to yellowish setae .............................................................45
- Abdomen covered with black setae ......................................................................47
45 Mesoscutum with four shiny vittae; wing deeply brown, broadly rounded (Fig. 40); fore coxa with fine setae anteriorly . . . . .
...................................................................... R. (P.) septentrionalis Sinclair et al.
- Mesoscutum uniformly pruinose; wing faintly brownish infuscate, of normal shape; fore coxa with strong blunt-tipped setae
anteriorly ...........................................................................................46
46 Hind tibia with dense dorsal and ventral fine setae at least as long as tibia width . . . . . . . . . . . . . .R. (P.) kjellmanii Holmgren
- Hind tibia with ventral setae at most half-length of tibia width . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. (P.) frigida Sinclair et al.
47 Halter with yellowish knob ................................................................R. (P.) hoeli Frey
- Halter brownish .....................................................................................48
48 Scutum almost shiny or with shiny vittae ..................................................................49
- Scutum densely, more or less uniformly pruinescent .........................................................51
49 Anepisternum pruinose .....................................R. (P.) subtenuiterfilata Shamshev & Sinclair sp. nov.
- Anepisternum shiny ..................................................................................50
50 Scutum faintly greyish pruinose, with two shiny median vittae (sometimes additional less distinct vitta present on supra-alar
space) .........................................................................R. (P.) ursinella Melander
- Scutum shiny, at most very faintly pruinose in front of scutellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. (P.) longestylata Frey
51 Anal lobe of wing darker than wing tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. (P.) filicauda Henriksen & Lundbeck
- Wing uniformly infuscate ..............................................................................52
52 Scutellum with 4 setae [see also couplet 47] ...................................................R. (P.) hoeli Frey
- Scutellum with 6–8 setae ..............................................................................53
53 Upper proepisternum in front of spiracle with 3 setae; no distinct basal costal seta (Note: this species is known after the
lectotype female only and the range of variability of these characters is unclear.) . . . . . . . . . . . . R. (P.) nordqvistii Holmgren
- Upper proepisternum in front of spiracle without setae; usually 1 more or less distinct short basal costal seta . . . . . . . . . . . . 54
54 Mesonotal setae stronger, bristle-like; presutural dorsocentrals biserial, separated from supra-alars by broad bare space . . . . .
..............................................................R. (P.) wrangeli Shamshev & Sinclair sp. nov.
- Mesonotal setae fine; presutural dorsocentrals 3–4-serial, barely separated from supra-alars (Fig. 68) . . . . . . . . . . . . . . . . . . . .
....................................................................................Rhamphomyia sp. 1
SHAMSHEV ET AL.
16 · Zootaxa 4848 (1) © 2020 Magnolia Press
Rhamphomyia (Ctenempis) albopilosa Coquillett
(Fig. 9)
Rhamphomyia albopilosa Coquillett, 1900: 418. Type-locality: Berg Bay, Alaska, USA.
Other references: Sinclair et al., 2019: 9 (lectotype designation, redescription, full list of references).
Rhamphomyia calvimontis Cockerell, 1916: 123. Type-locality: Baldy Mtn., Colorado, USA.
Other references: Sinclair et al., 2019: 9 (synonym, lectotype designation).
Rhamphomyia (Dasyrhamphomyia) wuorentausi Frey, 1922: 67. Type-locality: Dudinka, Krasnoyarskiy Territory, Russia.
Other references: Sinclair et al., 2019: 9 (synonym, lectotype designation).
Rhamphomyia (Ctenempis) wuorentausi: Frey, 1935: 3 (key and new subgenus); Frey, 1955b: 513 (revision).
Rhamphomyia (Rhamphomyia) wuorentausi: Chvála & Wagner, 1989: 292 (catalogue); Yang et al., 2007: 191 (catalogue);
Shamshev, 2016: 84 (checklist).
FIGURE 9. Rhamphomyia (Ctenempis) albopilosa Coquillett, male habitus, lateral view.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 17
Material examined. RUSSIA. Chukotka AO (Wrangel Island): Somnitelnaya Bay, valley of river Somnitelnaya,
Mineev Mtns, 150 m, shingle, on flower of Potentilla emarginata, 22.vii.1966, KBG (1 ♀, ZIN); same locality, on
inflorescence of Polygonum ovalifolium, 19.vii.1966, KBG (1 ♂, ZIN); same locality, on flower Dryas integrifolia,
19, 22.vii.1966, KBG (2 ♂, ZIN); 5 km N Somnitelnaya Bay, valley of river Somnitelnaya, foothills, 150 m,
25.vii.1972, KBG (7 ♂, 11 ♀, ZIN); near north slope of Berri Peak, 6 km ESE mountain Sovetskaya, meadow near
stream, 300 m, 12.vii.1972, KBG (1 ♂, ZIN); upper flow of river Khischniki, 7 km SE mountain Sovetskaya, Dryas
tundra near river, 200 m, 11.vii.1972, KBG (1 ♀, ZIN); same locality, meadow near stream, 200 m, 12.vii.1972,
KBG (1 ♂, ZIN); same locality, on Dryas, 14.vii.1972, KBG (1 ♂, 1 ♀, ZIN); same locality, 15.vii.1972, KBG (1
♂, 3 ♀, ZIN); basin of river Khischniki, 8 km SE mountain Sovetskaya, meadow near stream, 250 m, 13.vii.1972,
KBG (8 ♂, 16 ♀, ZIN); environs of mountain Tundrovaya, near stream, 18.vii.1972, KBG (13 ♂, 7 ♀, ZIN); same
locality, BT 3, PT, 1–19.vii.2015, LFV (1 ♀, ZIN); middle flow of river Mamontovaya, N Perkatkun, osier-bed in
river valley, 17,21,29.vii.1972, KBG (10 ♂, 1 ♀, ZIN); same locality, 71°10′N 179°45′W, BT 2’, 8.vii.2006, OAK
(1 ♂, 1 ♀, ZIN); same locality, BT 14, 9.vii.2006, OAK (1 ♀, ZIN); same locality and biotope, Sw, 2.vii.2015,
OAK (2 ♂, 1 ♀, ZIN); same locality, Sw, BT 20A, 1.vii.2015, OAK (1 ♀, ZIN); same locality and biotope, YPT,
2–4.vii.2015, OAK (2 ♂, ZIN); same locality and biotope, PT, 1–23.vii.2015, OAK (1 ♀, ZIN); same locality, Sw,
BT 1A, 2.vii.2015, OAK (1 ♀, ZIN); same locality, litter, BT 7, 1.vii.2015, OAK (1 ♂, ZIN); same locality, Sw,
BT 6, 2.vii.2015, OAK (1 ♂, ZIN); same locality, Sw, BT 11, 30.vi.2015, OAK (1 ♀, ZIN); same locality, Sw, BT
5, 30.vi.2015, OAK (1 ♀, ZIN); same locality, in house, 2.vii.2015, OAK (2 ♂, 1 ♀, ZIN); upper flow of river
Neizvestnaya, 71°13′N 179°19′W, BT 7А, 5.vii.2006, OAK (4 ♂, 5 ♀, ZIN); same locality, Sw, BT 4A, 22.vi.2015,
OAK (1 ♀, ZIN); spures of Pervaya Mtn, Sw, BT 11, 28.vi.2015, OAK (2 ♂, 1 ♀, ZIN); same locality, Sw, BT
13, 28.vi.2015, OAK (1 ♂, ZIN); middle flow of river Neizvestnaya, 71°20.287′N, 179°29.779′W, BT 1, YPT,
4–5.vii.2019, U.V. Babiy (1 ♂, ZIN).
Recognition. Large (wing length 6–7 mm) species with setose prosternum; thorax and abdomen covered with
long pale setae, legs long slender. Male (Fig. 9) holoptic; scutum mostly subshiny (dorsal view); mesonotal setae
mostly hair-like, 4–6 strong npl, at least 10 sctl, acr biserial, presutural dc multiserial; legs dark, fore basitarsus
distinctly thickened (slightly broader than tibia width at apex), with long setae (except dorsal and ventral face), hind
basitarsus usually slightly thickened but often quite slender, with long setae dorsally (more numerous and longer
in southern populations); wing infuscate, basal costal seta absent, CuA+CuP reaching wing margin but sometimes
weakened on about middle, halter with yellowish knob; abdomen with faintly pruinescent tergites dorsally; segments
6–8 modified, tergite 6 broadened, deeply concave posteriorly, sternite 6 inflated, with dense transverse cluster of
strong golden setae, sternite 7 with pair of horn-like projections ventrally. Female legs without pennate setae; wing
brownish, broadened.
Distribution. Holarctic; in North America, R. albopilosa is widespread across the low arctic, with isolated
southern records (Sinclair et al. 2019). In Eurasia, the species occurs in Eastern and Western Siberia (Shamshev
2016). Among the Russian Arctic islands, R. albopilosa is known only from Wrangel Island.
Habitat. On Wrangel Island this species is found in various biotopes.
Rhamphomyia (Dasyrhamphomyia) brusewitzii Holmgren
(Fig. 10)
Rhamphomyia brusewitzii Holmgren, 1880: 20. Type locality (by lectotype designation): Novaya Zemlya Islands, Arkhangelskaya
Province, Russia.
Other references: Holmgren & Aurivillius, 1883: 163 (expedition results); Shamshev & Sinclair, 2018: 308 (lectotype designation,
redescription).
Rhamphomyia brusewitzi: Frey, 1915: 11 (faunistic survey); 1922: 67 (key).
Rhamphomyia (Rhamphomyia) hovgaardii: Frey, 1955b: 516 (revision); Chvála & Wagner, 1989: 286 (catalogue); Yang et al.,
2007: 183 (catalogue); Shamshev, 2016: 76 (checklist).
Material examined. RUSSIA. Archangelskaya Prov. (Novaya Zemlya Archipelago): Beluzhja Bay, 5.vii.1904,
Sokolovsky (1 ♂, 1 ♀, ZIN); foot of Pila Mtn, Matochkin Shar, 9.viii.1901, Timofeev (1 ♂, ZIN); Matochkin Shar,
winter hut Razmyslova, 28.viii.1923, Lozina-Lozinsky (1 ♂, 1 ♀, ZIN); Matochkin Shar, brook Nochuev, 18–
19.vii.1925, Vakulenko (18 ♂, 10 ♀, ZIN). Krasnoyarskiy Terr. (Dikson Island): wet marsh in lowland between
hills (some specimens on Dryas punctata), 28–30.vii.1948, Korotkevich (6 ♂, 5 ♀, ZIN).
SHAMSHEV ET AL.
18 · Zootaxa 4848 (1) © 2020 Magnolia Press
Recognition. Large, blackish brown species (wing length 6–6.5 mm). Male (Fig. 10) eyes dichoptic, separated by
very narrow frons; thorax black setose, scutum densely brownish pruinescent, without distinct vittae; all mesonotal
setae strong, bristle-like, long, presutural dc 2–3-serial, acr biserial, 10–12 sctl; legs robust, entirely dark brown,
hind femur entirely setose, hind tibia slender; wing brownish infuscate, halter brown; cerci directed anteriorly, short,
prolonged nearly to middle of tergite 5, epandrium brownish to yellowish brown. Female legs not pennate.
Distribution. Eurasia. This species is known only from the Russian Arctic territories, including Novaya Zemlya
Archipelago, Taymyr Peninsula (Shamshev & Sinclair 2018) and Dikson Island.
Rhamphomyia (Dasyrhamphomyia) erinacioides Malloch
(Fig. 11)
Rhamphomyia erinacioides Malloch, 1919: 45. Type-locality: Camden Bay, Alaska, USA.
Rhamphomyia (Rhamphomyia) erinacioides: Shamshev, 2016: 75 (checklist).
Rhamphomyia (Dasyrhamphomyia) erinacioides: Sinclair et al., 2019: 14 (type material, redescription, full list of references).
Material examined. RUSSIA. Archangelskeya Prov. (Novaya Zemlya Archipelago): Beluzhya Bay, 5.vii.1904,
Sokolovskij (3 ♂, 1 ♀, ZIN); foot of Pila Mtn, Matochkin Shar, 9.viii.1901, Timofeev (1 ♂, 1 ♀, ZIN). Yakutia
(New Siberian Islands): centre of Kotelny Island, upper flow of river Balykhtakh near mouth of river Tuguttakh,
stream valley on flowers of Caltha, 15–16.vii.1973, KBG (34 ♂, 18 ♀, ZIN); same locality, arctic desert on flowers,
15.vii.1973, KBG (1 ♂, 1 ♀, ZIN). Chukotka AO (Wrangel Island): middle flow of Neozhidannaya River,
71°01′N 179°09′E, BT 1, YPT, 12–16.vii.2016, LFV (1 ♀, ZIN); same locality, BT 3, YPT, 12–16.vii.2016, LFV
(1 ♂, 8 ♀, ZIN); same locality, BT 4, YPT, 12–16.vii.2016, LFV (4 ♀, ZIN); same locality, BT A, Sw, 8.vii.2019,
OAK (1 ♀, ZIN); same locality, BT W, Sw, 8.vii.2019, OAK (1 ♂, ZIN); same locality, BT B, Sw, 8.vii.2019, OAK
(1 ♀, ZIN); same locality, BT G, Sw, 8–9.vii.2019, OAK (3 ♀, ZIN); 2 km W Somnitelnaya Bay, spotted tundra,
9.vii.1972, KBG (3 ♂, 6 ♀, ZIN); Somnitelnaya Bay, on flowers of Dryas integrifolia, 21–26.vii.1966, KBG (12
♂, 16 ♀, ZIN); near northern slope of Berri Peak, 6 km ESE Sovetskaya Mtn, 300 m, swamp on slope, 12.vii.1972,
KBG (1 ♂, ZIN); Sovetskaya Mtn, snowfield near top, 1000 m, 14.vii.1972, KBG (1 ♀, ZIN); SE part of island,
tundra, 2.viii.1931 & 26.vii.1933, А. Mineev (1 ♂, 1 ♀, ZIN); Rogers Bay, 20.vii.1932, А. Mineev (6 ♂, 4 ♀,
ZIN); same locality, 19.vii.1972, KBG (1 ♀, ZIN); same locality, 70°59′N 178°29′W, BT 1, YPT, 10–13.vii.2019,
OAK (1 ♂, 1 ♀, ZIN); same locality, Ushakovskoe vill., 70°59′N 178°30′W, BT A, YPT, 10–13.vii.2019, OAK (2
♀, ZIN); environs of Tundrovaya Mtn, tundra and near stream, 18.vii.1972, KBG (2 ♂, 1 ♀, ZIN); same locality,
71°18.469′N 179°44.327′W, BT 3, 5–15.vii.2018, U.V. Babiy (1 ♂, ZIN); same locality, 71°17.788′N 179°47.81′W,
BT 5, 15–26.vii.2018, U.V. Babiy (1 ♂, ZIN). USA. Alaska: Barrow, ex. Dryas, 18–26.vii.1955, sweeping arctic
tundra (13 ♂, 10 ♀, CNC); Inaru River, 25–28.vii.1956; ex. Polygonum and dandelion flowers (6 ♂, 1 ♀, CNC).
Recognition. Mid-sized blackish species (wing length about 5 mm). Male (Fig. 11) dichoptic, frons very broad;
thorax black setose, scutum densely brownish pruinescent, without distinct vittae; all mesonotal setae fine, hair-like,
long, presutural dc multiserial, acr 2–4-serial, at least 10 sctl; legs robust, entirely dark brown, all tibiae only pilose
ventrally, hind femur without setae on about apical 3/4 ventrally, only pale pilose, hind tibia very slender; wing
faintly infuscate, halter brown; cerci prolonged anteriorly, short, extended nearly to middle of tergite 5, epandrium
brown. Female legs not pennate.
Distribution. Holarctic species widely distributed across extreme northern Russia, including Novaya Zemlya
Islands, Taymyr Peninsula, New Siberian Islands and Wrangel Island. In North America, R. erinacioides is recorded
on Barter Island in Camden Bay, Alaska, the nearby mainland coast (Sinclair et al. 2019) and newly recorded here
from the region surrounding Barrow.
Habitat. On Wrangel Island, in the 1960s and 70s this species was most numerous on the coastal Southern
plane; in the mountains at around 1000 m. In the 2000s, the greatest abundance of this species was observed in areas
with more severe climatic conditions; e.g., in the northern variant of the Arctic tundra subzone (Neozhidannaya
River), where it was found in various habitats, including zonal spotted tundra. In the warmer central region of the
island, only a single specimen was captured.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 19
FIGURES 10–13. Rhamphomyia (Dasyrhamphomyia), male habitus, lateral view. 10. R. (D.) brusewitzii Holmgren; 11. R. (D.)
erinacioides Malloch; 12. R. (D.) hovgaardii Holmgren; 13. R. (D.) nigrita (Zetterstedt).
Rhamphomyia (Dasyrhamphomyia) gorodkovi Shamshev, Sinclair & Saigusa sp. nov.
(Figs 14–19)
urn:lsid:zoobank.org:act:34153559-3823-4932-A856-693999B0C7A4
Type material. HOLOTYPE ♂, labelled: [RUSSIA, Chukotka AO:] “[in Cyrillic, Russian] Apapelkhin, 15
km/ NOO Pevek, Chukot. [=Chukotka]/ Gorodkov 28.vi.[1]963”; “shrubby tundra”; “Holotypus/ Rhamphomyia/
gorodkovi Shamshev, Sinclair, Saigusa sp. nov. [red label]” (INS_DIP_0000613, ZIN).
PARATYPES: RUSSIA. Chukotka AO (Wrangel Island): upper flow of river Neizvestnaya, 71°13′N
179°19′W, BT 3G, 4.vii.2006, OAK (1 ♂, ZIN); same locality, 71°12.96′N, 179°20.09′W, 125 m, BT 12, Sw,
24.vi.2015, OAK (1 ♀, ZIN) same locality, 71°13.02′N, 179°19.508′W, 120 m, BT 3, Sw, 21.vi.2015, OAK (1 ♂,
ZIN). Chukotka mainland: same locality as holotype, 28.vi.1963, KBG (10 ♂, 6 ♀, ZIN; 2 ♂, 1 ♀, CNC); Iultin,
waterlogged sedge tundra, 21.vii.1963, KBG (3 ♂, 2 ♀, ZIN); Krasnoarmeiskiy, Chaunskiy District, osier-bed in
valley, 8.vii.1963, KBG (1 ♂, ZIN); 24 km SE Pevek, 150 m, osier-bed in valley, 1.vii.1963, KBG (1 ♂, ZIN);
valley of river Ichuvisi, Komsomolskiy mine, Chaunskiy District, shrubby tundra, 5.vii.1963, KBG (3 ♂, 2 ♀,
SHAMSHEV ET AL.
20 · Zootaxa 4848 (1) © 2020 Magnolia Press
ZIN); Komsomolskiy mine, Chaunskiy District, tundra, 4.vii.1963, KBG (17 ♂, 3 ♀, ZIN); Schmidt, southern slope,
tundra with willow, 18.vii.1963, KBG (1 ♀, ZIN); 20 km SSE Iultin, river Amguema, bush of Alnaster fruticosa,
22.vii.1963, KBG (3 ♂, ZIN); Schmidt, on snowfield, 16.vii.1966, KBG (1 ♂, ZIN); Schmidt, 5 km SW of village,
southern slope, tundra with willow, 11.vii.1971, KBG (3 ♂, 3 ♀, ZIN). Krasnoyarskiy Terr.: Taymyr, village
Agapa, Pyasina, moist shrubby tundra, 14.vii.1967, KBG (1 ♀, ZIN). Yakutia: Chokurdakh, Indigirka River, river
bank, slope, shrubby tundra, 12.vii.1966, KBG (1 ♀, ZIN); lower flow of Indigirka River, 71°N, border of tundra
and forest-tundra, 20.vi.1973, V. Flint (2 ♂, 1 ♀, ZMMU; 1 ♂, 1 ♀, CULSP). USA. Alaska: Cape Thompson,
29.vii.1960, H.E. Erdman (1 ♀, CNC).
Additional material. RUSSIA. Chukotka AO: Wrangel Island: middle flow of river Neozhidannay, 71°01′N
179°09′E, BT A, YPT, 8–9.vii.2019, OAK (1 ♂, ZIN); same locality, BT B, Sw, 8–9.vii.2019, OAK (1 ♂, ZIN);
same locality, BT W, YPT, 8–9.vii.2019, OAK (1 ♂, ZIN); same locality, BT G, 8–9.vii.2019, OAK (5 ♂, 4 ♀, ZIN);
same locality, YPT, BT 4, 12–16.vii.2016, LFV (2 ♂, 3 ♀, ZIN); same locality, YPT, BT 1, 12–16.vii.2016, LFV (1
♂, ZIN); same locality, YPT, BT 3, 12–16.vii.2016, LFV (2 ♂, 2 ♀, ZIN).
Diagnosis. A large species (body about 6.5 mm) with body and abdomen densely light grey pruinescent, halteres
yellow, acrostichals multiserial. Male: holoptic, legs uniformly brownish, hind femur with numerous short setae
over entire length ventrally, hind tibia thickened towards apex; wing whitish; cerci short, extended only to middle
of tergite 6. Female: legs extensively yellowish, with simple setae, hind tibia broadened toward apex, wings faintly
infuscate.
Description. Male (Fig. 14). Body length 6.1–6.4 (lectotype 6.3); wing length 6.0–6.2 (lectotype 6.2) mm. Head
with dense light greyish pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Holoptic,
eye with upper ommatidia enlarged. Frons represented by very small triangular space below ocellar tubercle and
larger subtriangular space above antennae, with scattered marginal setulae above antennae. Face parallel-sided,
bare. Ocellar triangle with several fine setae of different lengths. Occiput with numerous moderately long setae
laterally, including postoculars; postgena with numerous finer setae than occiput. Antenna with scape and pedicel
brownish, postpedicel somewhat darker, blackish brown; scape and pedicel short, scape slightly longer, both with
short setulae; postpedicel conical, 2.4–2.6 longer than wide; stylus short, 2.5–2.6 times shorter than postpedicel.
Palpus dark; with long, dark fine setae. Proboscis with labrum dark reddish brown, 1.2–1.3X head height.
Thorax dark in ground-colour, largely densely light grey pruinescent; with black setation. Scutum almost
uniformly light grey, with 2 indistinct, narrow, brownish grey vittae along dc rows (dorsal view), sometimes,
hardly visible vitta along acr rows present. Proepisternum with tuft of numerous long setae on lower section;
upper proepisternum in front of spiracle with 3–5 fine setae. Prosternum bare. Antepronotum with numerous short
setae. Postpronotal lobe with numerous almost uniform fine setae. Mesonotum mostly with fine setae, including
numerous undifferentiated similar presut spal, npl and psut spal (some posterior npl and psut spal somewhat longer
and stronger), 2–3 pal with several shorter setae (number and robustness variable); 6–10 of subequally long sctl
(sometimes with additional shorter setae); acr short, multiserial, absent on prescutellar depression; presutural dc
short and multiserial, postsutural dc less numerous, long and arranged in 2–3 irregular rows. Laterotergite with
numerous long fine setae. Anterior and posterior spiracles yellowish.
Legs almost uniformly brownish, only extreme base of tibiae somewhat yellowish, mostly subshiny, coxae
and trochanters with denser greyish pruinescence; black setose. Femora slender, of subequal width; mid tibia
slightly stouter than fore tibia, hind tibia evenly thickened toward apex; all tarsomeres slender. Mid and hind femora
whitish pilose ventrally (more distinctly on hind femur); fore and mid femora clothed in moderately long, thin
setae, mid femur with more numerous and denser anteroventral setae near base; mid femur with similar setation
but anteroventral setae more numerous and denser near base; hind femur entirely covered with setulae ventrally,
bearing complete rows of short setae anteroventrally (shorter than half of femur width). All tibiae covered with
moderately long, thin setae posterodorsally (at most as long as corresponding tibia width), without strong setae; no
seta in posteroapical comb of hind tibia. All tarsomeres covered with short, simple setae (except somewhat longer
setae of subapical circlet).
Wing membrane somewhat whitish, mostly with brownish veins; all veins complete (except Sc); CuA+CuP
(anal vein) mostly weakened but short subapical portion normally sclerotised. Pterostigma brownish yellow; basal
costa seta absent. Anal lobe well-developed; axillary incision acute but almost 90°. Squama yellow, dark fringed.
Halter yellow.
Abdomen dark, densely light grey pruinescent (concolorous with thorax); covered with black setae longer on
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 21
tergites laterally and shorter on tergites dorsally. Segments before segment 8 unmodified. Segment 8 with tergite
and sternite separated. Tergite 8 strongly upturned, laying almost vertically to body axis, with posterior margin
slightly recurved, with numerous setae along posterior margin. Sternite 8 simple, scoop-like, with numerous long
setae along posterior margin.
FIGURES 14–19. Rhamphomyia (Dasyrhamphomyia) gorodkovi sp. nov. 14. holotype male, habitus, lateral view; 15. male
postabdomen, holotype, lateral view; 16. male terminalia, lateral view; 17. male terminalia, dorsal view; 18. phallus, lateral
view; 19. female habitus, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium; goncx
apod –gonocoxal apodeme; hypd—hypandrium; ph—phallus.
SHAMSHEV ET AL.
22 · Zootaxa 4848 (1) © 2020 Magnolia Press
Terminalia (Figs 15–18). Cerci yellowish; fused to epandrium (but suture distinct), broadly concave and
flattened medially, strongly prolonged anteriorly, forming small subglobular window-like space subapically (dorsal
view), without pair of subepandrial lobes beneath; apical part of cercus (before epandrium) short, extended only
to tergite 6, subtriangular viewed laterally, rounded apically, covered with dark setulae; posterior part of cercus
broadly digitiform, rounded apically, only slightly extended beyond epandrium, with numerous dark spinule-like
setulae ventrally. Epandrium usually brownish, sometimes yellowish or reddish brown, faintly greyish pruinescent,
mostly with short dark fine setae longer on posterior projection; rather subglobular (lateral view), with short,
rounded apically projection on lower part posteriorly. Hypandrium yellowish, subtriangular (ventral view), entirely
sclerotized, bare; gonocoxal apodeme directed posteriorly, large, broadened apically. Phallus (Fig. 18) yellowish;
short, mostly hidden (lateral view); gently arched, broad on about basal half, becoming evenly slenderer beyond
epandrial projection; with two minute lateral “spinules” on apical part. Ejaculatory apodeme extended far beyond
basal curvature of phallus, with narrow lateral wings and very broad vertical wing.
Female (Fig. 19). Body length 6.4; wing length 5.9 mm. Similar to male, except eyes dichoptic, ommatidia of
equal size; frons broad, somewhat widened toward ocellar tubercle; mesonotum, legs and abdomen with distinctly
shorter setation; legs paler, largely rather brownish yellow, with simple setae (sometimes posteroventral setae on
mid and hind femora slightly flattened); wing membrane faintly brownish infuscate; abdominal tergite 8 pointed
posteriorly; cercus black, long, slender, covered with dark setulae.
Distribution. Holarctic; in Eurasia, the new species is broadly distributed across subarctic areas of Russia
(from Taymyr to Chukotka). Among the Russian Arctic islands the species is known only from Wrangel Island. In
North America, this species is known from Alaska and Yukon, but unknown from Canadian Arctic islands.
Etymology. The specific name is a patronym in honour of the late Kirill Borisovich Gorodkov, Russian Dipterist
(ZIN, St. Petersburg), who collected many of the specimens used in this study, as well as sorted and identified many
Palaearctic Rhamphomyia housed in the ZIN.
Remarks. The new species resembles strongly the Holarctic species R. villipes Coquillett, 1900 (known for
a long time in Eurasia as R. hambergi Frey). However, R. gorodkovi sp. nov. can be readily distinguished from R.
villipes primarily by the whitish wings of the male (hyaline to very faintly infuscate in R. villipes) and simple setose
mid and hind legs of the female (mid, hind femora and hind tibia with pennate setae in R. villipes). In addition, in the
male of R. villipes the legs are covered with long hair-like setae, fore and mid tibiae somewhat thickened and mid
femur slightly arcuate. In the Chukotka Peninsula, both species appear to occur as sympatric populations. Among
the material from North America, this species was recognised as Rhamphomyia (Dasyrhamphomyia) sp. 8 (Saigusa
unpubl. data).
Rhamphomyia (Dasyrhamphomyia) hovgaardii Holmgren
(Fig. 12)
Rhamphomyia hovgaardii Holmgren, 1880: 21. Type locality (by lectotype designation): Novaya Zemlya Islands, Arkhangelskaya
Province, Russia.
Other references: Holmgren & Aurivillius, 1883: 162; Melander, 1928: 195 (catalogue); Gorodkov & Kovalev, 1969: 629
(key).
Rhamphomyia zaitzevi Becker, 1915: 54. Type-locality (by lectotype designation): upper reaches of the Kara River (about
68°39′N 65°52′E), Yamalo-Nenets, Russia.
Other references: Sinclair et al., 2019: 17 (synonym, lectotype designation).
Rhamphomyia (Dasyrhamphomyia) hovgaardi: Frey, 1922: 66 (key); Frey, 1955b: 516 (revision).
Rhamphomyia (Rhamphomyia) hovgaardi: Chvála & Wagner, 1989: 285 (catalogue); Shamshev, 2001b: 334 (key); Yang et al.,
2007: 183 (catalogue).
Rhamphomyia (Rhamphomyia) hovgaardii: Shamshev, 2016: 76 (checklist).
Rhamphomyia (Dasyrhamphomyia) hovgaardii: Shamshev & Sinclair, 2018: 310 (revision, lectotype designated); Sinclair et
al., 2019: 17 (redescription).
Rhamphomyia (Rhamphomyia) zaitzevi: Shamshev, 2001b: 334 (key).
Rhamphomyia zaitsevi. Incorrect subsequent spelling of zaitzevi Becker, 1915 (Chvála & Wagner, 1989: 292; Yang et al., 2007:
191; Shamshev, 2016: 84).
Material examined. RUSSIA. Krasnoyarskiy Terr. (Dikson Island): small island “Konus”, 31.vii.1948,
Korotkevich (1 ♂, ZIN); marsh in lowland between hills, 30.vii.1948, Korotkevich (1 ♀, ZIN). Chukotka AO
(Wrangel Island): middle flow of Neozhidannaya River, 71°01′N 179°09′E, BT 4, YPT, 12–16.vii.2016, LFV (1
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 23
♀, ZIN); environs of Tundrovaya Mtn, tundra nesting-place of gooses, 18.vii.1972, KBG (1 ♀, ZIN); middle flow
of Mamontovaya River, N Perkatkun, waterlogged coast, 17.vii.1972, KBG (2 ♂, 3 ♀, ZIN); same locality, small
swamp, 29.vii.1972, KBG (1 ♂, 3 ♀, ZIN); same locality, 71°10′N 179°45′W, BT 20, 12.vii.2006, OAK (3 ♀,
ZIN); same locality and biotope, Sw, 1.vii.2015, OAK (2 ♂, 3 ♀, ZIN); upper flow of Neizvestnaya River, 71°13′N
179°19′W, BT 12, PT, 24.vi.–6.vii.2015, OAK (1 ♀, ZIN).
Recognition. Large, blackish brown species (wing length 6.3–7 mm). Male (Fig. 12) eyes holoptic; thorax
black setose, scutum densely brownish pruinescent, without distinct vittae; almost all mesonotal setae fine, hair-like
(npl usually slightly stronger), long, presutural dc 3–5-serial, acr 2–3-serial; legs robust, entirely dark brown, hind
femur entirely setose, hind tibia slender; wing brownish infuscate, halter brown; cerci directed anteriorly, short,
prolonged nearly to middle of tergite 5, epandrium brownish. Female wing slightly darker and broader; mid femur
with pennate posteroventral setae, hind femur with pennate anterodorsal and posteroventral setae, hind tibia with
pennate setae dorsally and ventrally (on about middle).
Distribution. Holmgren (1880) described R. hovgaardii from Novaya Zemlya, however, later he noted also
Vaygach Island (Holmgren & Aurivillius 1883; see Shamshev & Sinclair 2018). Holarctic species; in Eurasia, R.
hovgaardii extends across arctic and subarctic Russia, including both continental areas (Taymyr, Yakutia, Chukotka)
and islands (Novaya Zemlya, Vaygach, Wrangel). In North America, this species ranges in the low arctic from the
western edge of Hudson Bay to Alaska (Sinclair et al. 2019).
Habitat. On Wrangel Island, this species was collected mainly in wet habitats.
Rhamphomyia (Dasyrhamphomyia) mallochi Shamshev, Sinclair & Saigusa sp. nov.
(Figs 20–25)
urn:lsid:zoobank.org:act:26E7A46E-1A47-4193-A5FF-FA5618C8E98E
Type material. HOLOTYPE ♂, labelled: [RUSSIA, Chukotka AO:] “[in Cyrillic, Russian] Pevek [69°42′N
170°19′N], Chukotskij nats. [=natsionalny, national] okr. [=okrug, name of administrative unit] archadovaya/
tundra./ Gorodkov, 28.vi.[1]963”; “Holotypus/ Rhamphomyia/ mallochi/ Shamshev, Sinclair, Saigusa sp. nov. [red
label]” (INS_DIP_0000614, ZIN).
PARATYPES: CANADA. Yukon: Dawson, 10,11.vii.1949, W.W. Judd (3 ♀, CNC); km 465 Dempster
Hwy, 23–25.vi.1980, D.M. Wood & D. Lafontaine (1 ♂, CNC); Richardson Mts, 66°26.5′N 135°48′W, 3000 ft,
8.vii.1982, D. Wood (2 ♂, CNC). RUSSIA. Chukotka AO (Wrangel Island): SE part of Rogers Bay, 20.vii.1932,
A. Mineev (1 ♀, ZIN); SE part of Rogers Bay, 24.vii.1932, A. Mineev (1 ♀, ZIN); environs of Somnitelnaya Bay,
valley of river Somnitelnaya, Mineev Mtns, 150 m, shingle, on flower of Dryas integrifolia, 19.vii.1966, KBG
(1 ♂, ZIN); lower flow of river Khischniki, foothills of Mineev Mtns, marshy tundra, 11.vii.1972, KBG (1 ♂,
1 ♀, CNC); middle flow of river Neozhidannay, 71°01′N 179°09′E, BT G, 8–9.vii.2019, OAK (3 ♂, 1 ♀, ZIN);
same locality, BT A, YPT, 8–9.vii.2019, OAK (2 ♀, ZIN); same locality, BT W, YPT, 8–9.vii.2019, OAK (1 ♂,
ZIN); same locality, BT D, YPT, 8–9.vii.2019, OAK (1 ♀, ZIN); Inkali Mtn, 71°05′N 179°42′W, damp tundra,
Sw, 17.vii.2019, OAK (1 ♀, ZIN). Chukotka AO (Kolyuchin Island): Kolyuchin Island, 25–27.vii.1938, Hecker
(5 ♂, 5 ♀, ZIN; 1 ♂, 1 ♀, CULSP). Chukotka mainland: Apuka, island in delta of river Apuka, on flowers
of cloudberry Rubus chamaemorus, 2.vii.1959 KBG (1 ♂, ZIN); Apuka, Pakhachinskij Range, mountain tundra,
7.vii.1959, KBG (1 ♂, ZIN); Pevek, 200–300 m, moist tundra, 29.vi.1963, KBG (1 ♂, ZIN); same locality, dryad
tundra, 3.vii.1963, KBG (2 ♀, ZIN); same locality, osier-bed in valley, 8.vii.1963, KBG (1 ♂, ZIN); same locality,
moist sedge tundra, 11.vii.1963, KBG (1 ♂, ZIN); same locality, mossy-sedge tundra, 18.vii.1963, KBG (7 ♂, 10
♀, ZIN); Schmidt, on snow, 9.viii.1966, KBG (5 ♂, 1 ♀, ZIN); same locality, on snowfield, 16.vii.1966, KBG (6
♂, 7 ♀, ZIN); same locality, polygon tundra, 11.vii.1971, KBG (2 ♂, 1 ♀, ZIN); same locality, spotted tundra,
17.vii.1971, KBG (9 ♂, 5 ♀, ZIN); same locality, S slope tundra with willow, 11.vii.1971, KBG (1 ♀, ZIN); valley
of river Ichuvisi, Komsomolskij mine, Chaunskij District, shrubby tundra, 5.vii.1963, KBG (3 ♂, 2 ♀, ZIN); same
locality, sedge tundra and osier-bed in river valley, 4.vii.1963, KBG (10 ♂, 3 ♀, ZIN); Iultin, water-logged sedge
tundra, 21.vii.1963, KBG (11 ♂, 2 ♀, ZIN); 5 km N of Egvekinot, 27.vii.1963, KBG (1 ♂, ZIN). USA. Alaska:
Cape Thompson, 29.vii.1960, H.E. Erdman (1 ♀, CNC); Schrader Lake, 3–5.vii.1972, K.W. Philip (2 ♂, CNC).
Additional material: RUSSIA. Chukotka AO (Wrangel Island): middle flow of river Neozhidannaya,
71°01′N 179°09′E, YPT, BT 4, 12–16.vii.2016, LFV (1 ♂, ZIN); same locality, YPT, BT 3, 12–16.vii.2016, LFV
(3 ♀, ZIN).
SHAMSHEV ET AL.
24 · Zootaxa 4848 (1) © 2020 Magnolia Press
Diagnosis. A large species (body about 7 mm) with body and abdomen densely brownish pruinescent, halteres
yellow, acrostichals biserial. Male: eyes separated by very narrow frons, legs yellowish to brownish yellow, hind femur
with numerous short setae over entire length ventrally, hind tibia slender; wing membrane slightly infuscate; terminalia
yellowish, cerci extended only to tergite 5. Female: mid and hind femora with posteroventral pennate setae.
Description. Male (Fig. 20). Body length 6.9–7.3 (lectotype 7.1); wing length 6.9–7.4 (lectotype 7.3) mm.
Head with dense brownish pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Eyes
separated by frons; upper ommatidia slightly enlarged. Frons very narrow on middle part, at most as broad as
anterior ocellus, slightly broader just below ocellar tubercle, forming larger subtriangular space above antennae,
with scattered marginal setulae. Face broad, bare. Ocellar triangle with several fine setae of different lengths.
Occiput with numerous long, thin setae, including postoculars; postgena with numerous finer setae than occiput.
Antenna blackish brown; scape and pedicel short, scape slightly longer, both with short setulae; postpedicel conical,
nearly 3X longer than wide; stylus rather long, nearly 1.5X shorter than postpedicel. Palpus dark; with long, dark
fine setae. Proboscis with labrum dark reddish-brown, 1.2–1.3X head height.
Thorax dark in ground-colour, densely brownish pruinescent; with black setation; scutum brownish, without
vittae. Proepisternum with tuft of numerous long setae on lower section; upper proepisternum in front of spiracle
with 3–5 fine setae. Prosternum bare. Antepronotum with numerous short setae. Postpronotal lobe with numerous
almost uniform fine setae of different lengths, sometimes 1–2 setae somewhat stronger. Mesonotum mostly with fine
setae, including numerous undifferentiated similar presut spal, npl and psut spal (some posterior npl and psut spal
somewhat longer and stronger but their number and position are very variable, often even on right and left sides),
2–3 pal with several shorter setae (number and robustness variable); 8–10 of subequally long sctl and some number
of additional finer setae (number and robustness very variable); acr rather long, thin, arranged in 2 irregular rows,
absent on prescutellar depression; presutural dc long, arranged in 3–4 irregular rows, postsutural dc less numerous,
longer. Laterotergite with numerous long fine setae. Anterior and posterior spiracles yellowish.
Legs with coxae, trochanters and tarsi (except basal part of hind basitarsus) brownish, fore femur, mid femur
ventrally, fore tibia toward apex, mid and hind tibiae at apex brownish yellow, remaining parts yellowish; mostly
subshiny, coxae and trochanters with denser greyish pruinescence; black setose. Femora, tibiae and tarsomeres
slender; hind femur of subequal thickness to fore and mid femora. Mid and hind femora whitish pilose ventrally (more
distinctly on hind femur); fore femur with rather thin, moderately long setae anteroventrally and posteroventrally,
longest setae closer to base slightly longer than half of femur basal width; mid femur with numerous strong, short
setae anteroventrally and posteroventrally; hind femur entirely covered with setae ventrally (at most as long as half
of femur width). All tibiae with some short ad and pd setae stronger on hind tibia; no seta in posteroapical comb of
hind tibia. All tarsomeres covered with short, simple setae (except somewhat longer setae of subapical circlet); mid
and hind basitarsi with short, strong setae ventrally.
Wing membrane faintly infuscate, mostly with brownish veins; all veins complete (except Sc). Pterostigma
brownish yellow; basal costa seta absent. Anal lobe well-developed; axillary incision acute but closer to 90°. Squama
yellow, dark fringed. Halter yellow.
Abdomen dark, densely brownish pruinescent; covered with uniform black setae shorter on tergites dorsally.
Segments before segment 7 unmodified. Tergite 7 broadly concave posteriorly. Segment 8 with tergite and sternite
separated. Tergite 8 upturned, with posterior margin recurved anteriorly, with numerous setae along posterior margin.
Sternite 8 simple, scoop-like, with numerous long setae along posterior margin.
Terminalia (Figs 21–24). Cerci yellowish; fused to epandrium (but suture distinct), broadly concave and
flattened medially, strongly prolonged anteriorly, without pair of subepandrial lobes beneath; apical part of cercus
(before epandrium) short, extended to tergite 5, subrectangular viewed laterally, subtriangular, with straight inner
margin viewed dorsally, mostly bare, with some dark scattered setulae dorsally; posterior part of cercus very short,
rounded, not extended beyond epandrium, covered with fine setulae. Epandrium almost entirely yellowish, only
posterior digitiform projection brownish yellow (very rarely epandrium dark, 1 ♂ from Wrangel Island), faintly
greyish pruinescent, with moderately long dark setae on upper part and longer setae along lower margin, posterior
projection covered with black spinules dorsally; rather subrectangular (lateral view), somewhat humped dorsally,
with short, digitiform projection on lower part posteriorly. Hypandrium yellowish, subtriangular (ventral view),
entirely sclerotized, bare; gonocoxal apodeme large, broadened apically. Phallus (Fig. 24) yellowish; short, mostly
hidden (lateral view); gently arched, thick on about basal half, becoming abruptly slenderer beyond epandrial
projection; with two minute lateral “spinules” slightly beyond thickened portion. Ejaculatory apodeme extended far
beyond basal curvature of phallus, with narrow lateral wings and very broad vertical wing.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 25
Female (Fig. 25). Frons broad, parallel-sided; ocellar tubercle and occiput with somewhat shorter and
stronger setae. Thorax with greyish pruinescence, scutum with 3 indistinct brownish vittae along rows of acr and
dc; mesonotum with shorter setae. Legs often somewhat paler than in male (except coxae and trochanters) with
only fore femur and tarsomeres darkened; mid and hind femora with complete rows of moderately long, pennate
posteroventral setae. Abdomen with shorter setae; cercus long, slender, with dark setulae.
Distribution. Holarctic; in Eurasia, the new species is known from Chukotka mainland and nearby Arctic
islands (Kolyuchin, Wrangel); in North America, this species is known from Alaska and Yukon, but unknown from
the Canadian Arctic islands.
FIGURES 20–25. Rhamphomyia (Dasyrhamphomyia) mallochi sp. nov. 20. holotype male, habitus, lateral view; 21. male
postabdomen, holotype, dorso-lateral view; 22. male terminalia, lateral view; 23. male terminalia, dorsal view; 24. phallus,
lateral view; 25. female habitus, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium;
goncx apod—gonocoxal apodeme; hypd—hypandrium; ph—phallus.
SHAMSHEV ET AL.
26 · Zootaxa 4848 (1) © 2020 Magnolia Press
Etymology. The specific name is a patronym in honour of the J.R. Malloch, a world-renowned dipterist in the
first half of the 20th century, and among his numerous contributions were publications on Arctic Diptera (including
Empididae) of North America.
Remarks. The new species is most similar to R. hovgaardii differing from the latter primarily by brownish grey
pruinescent thorax and abdomen (dark brown in R. hovgaardii), yellowish halteres, extensively yellowish legs and
yellowish epandrium (brown in R. hovgaardii). The female of R. hovgaardii has pennate setae on the hind tibia. In
addition, the new species could be compared with R. plumipes (Meigen) and R. vesiculosa (Fallén). However, both
these species can be readily distiguished from R. mallochi sp. nov. primarily by the much longer cerci of the male
terminalia extending to abdominal tergite 2 and dark halteres.
Among material from North America, the species was recognised as Rhamphomyia (Dasyrhamphomyia) sp. 12
(Saigusa unpubl. data).
Habitat. Judging by the material of K.B. Gorodokov from 1950s–1970s, this species is most numerous in
coastal areas in northern Chukotka (Pevek, Schmidt), where it was recorded in various habitats. On Wrangel Island,
the largest series of specimens was collected in the colder regions of the island (Neozhidannaya River), where it was
most numerous in the zonal spotted tundra. Moreover, in the warmer central region, the only specimen was found
in a wetland below a snow field.
Rhamphomyia (Dasyrhamphomyia) nigrita (Zetterstedt)
(Fig. 13)
Rhamphomyza nigrita Zetterstedt, 1838: 567. Type-locality: Greenland.
Other references: Barták & Danielson, 2007: 107 (type data); Sinclair et al., 2019: 24 (full list of references).
R. (Dasyrhamphomyia) nigrita: Frey, 1922: 66 (key); Frey, 1955b: 518 (revision); Sinclair et al., 2019: 24 (Nearctic faunal
survey, redescription).
R. (Rhamphomyia) nigrita: Chvála & Wagner, 1989: 288 (catalogue); Shamshev, 2001b: 333 (key); Yang et al., 2007: 186
(catalogue); Barták, 2015: 576 (Greenland fauna); Shamshev, 2016: 78 (checklist).
Empis borealis Fabricius, 1780: 211 (junior primary homonym of Empis borealis Linnaeus, 1758). Type-locality: Greenland.
Rhamphomyia conservativa Malloch, 1919: 48. Type-locality: Bernard Harbour, Nunavut, Canada.
Material examined. RUSSIA. Chukotka AO (Wrangel Island): middle flow of Neozhidannaya River, 71°01′N
179°09′E, BT 2, YPT, 12–16.vii.2016, LFV (1 ♂, ZIN); environs of Somnitelnaya Bay, 6.vii.1972, KBG (1 ♀, ZIN);
4 km N of Somnitelnaya Bay, Somnitelnaya River, shore slope and flood-lands, 28.vii.1972, KBG (2 ♂, 1 ♀, ZIN);
5 km N of Somnitelnaya Bay, valley of Somnitelnaya River, foothills 150 m, on flowers of Dryas, 25, 28.vii.1972,
KBG (2 ♀, ZIN); Somnitelnaya Bay, valley of Somnitelnaya River, Mineev Mtns, 150 m, pebbles, on flower of
Potentilla emarginata, 22.vii.1966, KBG (1 ♀, ZIN); same locality, south slope of Mineev Mtns, 100 m, pebbles,
rubbly placer, 26.vii.1966, KBG (1 ♀, ZIN); same locality, Sw on Epilobium latifolium L., 23.vii.1971, KBG (1 ♀,
ZIN); basin of Khischniki River, 8 km SE of Sovetskaya Mtn, lawn near stream, 250 m, 13.vii.1972, KBG (1 ♂,
ZIN); environs of Tundrovaya Mtn, near stream, 18.vii.1972, KBG (1 ♀, ZIN); same locality, tundra, 18.vii.1972,
KBG (2 ♂, 2 ♀, ZIN); same locality, 71°18′N, 179°48′W, BT 3, PT, 1–19.vii.2015, LFV (1 ♀, ZIN); same locality,
BT 5, 1–19.vii.2015, PT, LFV (1 ♀, ZIN); same locality, BT 1, 71°18.342′N 179°48.367′W, 30.vi.–6.vii.2018, U.V.
Babiy (1 ♀, ZIN); middle flow of Mamontovaya River, N Perkatkun, osier-bed in river valley, 17, 21.vii.1972, KBG
(4 ♂, 2 ♀, ZIN); same locality and collector, 21.vii.1972 (19 ♂, 11 ♀, ZIN); same locality and collector, 29.vii.1972
(1 ♂, ZIN); same locality and collector, 18.vii.1972 (1 ♂, 2 ♀, ZIN); same locality, BT 4, 71°09′N 179°45′W, 26.vi.–
19.vii.2011, А.А. Rodionov (1 ♀, ZIN); same locality, 71°10′N 179°45′W, in house, 29.vi., 1.vii. 2015, OAK (2 ♀,
ZIN); same locality, BT 20A, Sw, 1–2.vii.2015, OAK (2 ♂, 3 ♀, ZIN); same locality, BT 14, Sw, 2.vii.2015, OAK
(7 ♂, 19 ♀, ZIN); same locality, BT 3, Sw, 1.vii.2015, OAK (1 ♂, 4 ♀, ZIN); same locality, BT 2, Sw, 2.vii.2015,
OAK (2 ♂, ZIN); same locality, BT 1A, Sw, 2.vii.2015, OAK (1 ♂, 1 ♀, ZIN); same locality and biotope, YPT,
2–4.vii.2015, OAK (1 ♂, 1 ♀, ZIN); same locality, BT 7, Sw, 1.vii.2015, OAK (1 ♂, 6 ♀, ZIN); same locality, BT 6,
Sw, 2–3.vii.2015, OAK (5 ♂, 5 ♀, ZIN); same locality, BT 6A, Sw, 1.vii.2015, OAK (1 ♀, ZIN); same locality, BT
11, Sw, 30.vi.2015, OAK (1 ♀, ZIN); same locality, BT 5, Sw, 30.vi.2015, OAK (2 ♀, ZIN); same locality, BT 10,
Sw, 30.vi.2015, OAK (1 ♀, ZIN); same locality, BT 17, Sw, 3.vii.2015, OAK (9 ♀, ZIN); same locality, Arctic fox
burrow, Sw, 1.vii.2015, OAK (1 ♂, 1 ♀, ZIN); same locality and biotope, PT, 1–23.vii.2015, OAK (1 ♀, ZIN); same
locality and biotope, Sw, 18.vii.2019, OAK (1 ♀, ZIN); brook Vesely, 71°11′N, 179°43′W, BT A, Sw, 18.vii.2019,
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 27
OAK (1 ♂, ZIN); spurs of Pervaya Mtn, 71°09′N, 179°27′W, BT 11, Sw, 28.vi.2015, OAK (5 ♂, 3 ♀, ZIN); same
locality and biotope, Sw, 13.vii.2015, OAK (1 ♀, ZIN); upper flow of Neizvestnaya River, 71°13′N, 179°19′W,
BT 4A, Sw 5.vii.2006, OAK (11 ex., SZM); same locality, BT 7A, 5.vii.2006, OAK (1 ex., SZM); same locality,
BT 3A, 6.vii.2006, OAK (1 ex., SZM); same locality, BT 12, Sw, 6.vii.2015, OAK (3 ♂, 2 ♀, ZIN); same locality,
BT 5A, PT, 26.vi.–10.vii.2015, OAK (1 ♀, ZIN); same locality and biotope, Sw, 12.vii.2015, OAK (1 ♀, ZIN);
same locality, BT 3B, Sw, 7.vii.2015, OAK (1 ♀, ZIN); same locality, BT C1, Sw, 21.vi.2015, OAK (3 ♀, ZIN);
same locality, wormwoods near house, Sw, 24.vi.2015, OAK (1 ♂, ZIN); middle flow of Neizvestnaya River, BT
1, YPT, 71°20.287′N 179°29.779′W, 5–14.vii.2018, U.V. Babiy (1 ♀, ZIN); same locality, BT 2, YPT, 71°20.416′N
179°29.651′W, 13–14.vii.2018, U.V. Babiy (2 ♀, ZIN).
Recognition. Large (wing length 6–7 mm) brownish grey species. Male (Fig. 13) holoptic; thorax black setose,
scutum densely brownish grey pruinescent, with dark vittae beneath acrostichal and dorsocentral rows; mesonotal
setae mostly fine, long, presutural dc and acr multiserial; legs robust, entirely dark brown, hind femur without setae
on about apical 2/3 ventrally, only densely pale pilose, hind tibia evenly thickened toward apex; wing brownish
infuscate, halter yellowish; cerci directed anteriorly, very long, extended nearly to middle of abdominal tergite 3,
epandrium dark. Female wing broadened; legs without pennate setae.
Distribution. A Holarctic species with huge area of distribution; in Eurasia, R. nigrita is widespread across
the low and high arctic of Norway, Sweden and Russia (with some records from the boreal zone; e.g., Altay,
Magadanskaya Province); in North America, from Greenland to Alaska and as far south as Churchill, Manitoba and
northern British Columbia (Sinclair et al. 2019). However, among the Russia Arctic islands, R. nigrita was found
only from Wrangel Island.
Habitat. On Wrangel Island this species is most common in the warmer central region of the island, where it
is found in various biotopes. In colder areas (Neozhidannaya River, coastal plain in Somnitelnaya Bay), a single
specimen was found.
Rhamphomyia (Eorhamphomyia) shewelli Sinclair, Vajda, Saigusa & Shamshev
(Fig. 26)
Rhamphomyia (Eorhamphomyia) shewelli Sinclair et al., 2019: 29. Type locality: Coral Harbour, Southampton Island,
Canada.
Material examined. RUSSIA. Chukotka AO (Wrangel Island): middle flow of Mamontovaya River, N
of Perkatkun, osier-bed in river valley, 17.vii.1972, KBG (1 ♀, ZIN); same locality, 71°10′N 179°45′W, BT 3,
29.vi.–9.vii.2006, OAK (1 ♂, ZIN); same locality and biotope, 9.vii.2006, OAK (1 ♀, ZIN); same locality and
biotope, 1.vii.2015, leg. OAK (3 ♀, ZIN); same locality, BT 1А, 2.vii.2015, OAK (1 ♂, ZIN); same locality,
BT 20А, 2–4.vii.2015, OAK (2 ♂, ZIN); same locality, 71°10′N 179°45′W, BT 7, 1.vii.2015, OAK (1 ♂, ZIN);
upper flow of river Neizvestnaya, 71°13′N 179°19′W, BT С1, 24.vi.2015, OAK (1 ♀, ZIN); same locality, BT 7А,
25.vi.–10.vii.2015, OAK (1 ♀, ZIN); middle flow of Neizvestnaya River, 71°20.287′N 179°29.779′W, BT 1, YPT,
5–14.vii.2018, U.V. Babiy (1 ♀, ZIN); same locality, 71°20.416′N 179°29.651′W, BT 2, YPT, 13–14.vii.2018, U.V.
Babiy (3 ♂, 2 ♀, ZIN); same locality and biotope, YPT, 4–5.vii.2019, U.V. Babiy (4 ♂, 4 ♀, ZIN).
Recognition. Mid-sized (wing length 4.7–5.3 mm) blackish species. Male (Fig. 26) holoptic; thorax black
setose, scutum with pair of pruinescent vittae anteriorly, sometimes lustrous posteriorly; supra-alar region with
broad shiny stripe, mesonotal setae well differentiated, presutural dc 3–4-serial, acr biserial, 6–8 sctl (rarely 10);
legs slender, entirely dark brown, no modified podomeres, hind femur with minute anteroventral setulae (except
several longer setae near extreme base), all tibiae with rows of long ad and pd setae; wing faintly infuscate, axillary
incision very acute, at most 45°, halter yellow; abdomen pruinose dorsally and narrowly shiny laterally; phallus
short, almost entirely hidden within epandrium and cerci, at most very short portion between lower margin of
epandrium and hypandrium visible, hypandrium well exposed, subtriangular. Female with somewhat darker wing,
legs without pennate setae. Besides the species noted in the key, R. shewelli should be compared primarily with R.
dorsata Becker known from Arctic continental territories of Eurasia. However, in the latter species the scutum is
broadly shiny laterally leaving only a very narrow greyish vitta beneath the acrostichals and the abdomen is entirely
shiny in dorsal view (Sinclair et al. 2019).
Distribution. Holarctic; in North America, this low arctic species is widespread across western and central
arctic region (Sinclair et al. 2019); in Eurasia, it is known only from Wrangel Island.
SHAMSHEV ET AL.
28 · Zootaxa 4848 (1) © 2020 Magnolia Press
Habitat. On Wrangel Island this species was collected only in the warmer central part of the island. The
majority of specimens were found in wet habitats with willow bushes, located in river valleys.
Rhamphomyia (Lundstroemiella) hybotina (Zetterstedt)
(Fig. 27)
Rhamphomyza hybotina Zetterstedt, 1838: 571. Type locality (by lectotype designation): Hacksten (Sweden).
Other references: Barták, 1999: 99 (type data, lectotype designation).
Rhamphomyza tenuicornis Zetterstedt, 1838: 571. Type locality (by lectotype designation): Hacksten (Sweden).
Other references: Barták, 1999: 100 (type data, lectotype designation).
Rhamphomyia (Lundstroemiella) hybotina Zetterstedt: Frey, 1922: 4 (key); 1955a: 434 (revision); Collin, 1961: 441 (revision);
Gorodkov & Kovalev, 1969: 613 (key); Barták, 1981: 451 (revision); 1985: 28 (revision); Chvála & Wagner, 1989: 310
(catalogue); Barták, 1999: 99 (type data); Shamshev, 2001b: 320 (key); Barták, 2006: 508 (key); 2007: 103 (catalogue);
Yang et al., 2007: 157 (catalogue); Shamshev, 2016: 56 (checklist).
Material examined. RUSSIA. Murmanskaya Prov. (Kildin Island): 13.vii.1908, L. Bianki (1 ♀, ZIN).
FIGURES 26, 27. Rhamphomyia. 26. R. (Eorhamphomyia) shewelli Sinclair, Vajda, Saigusa & Shamshev; 27. R.
(Lundstroemiella) hybotina (Zetterstedt).
Recognition. Small, blackish, pale-legged, delicate flies (body length about 3 mm). Male (Fig. 27) dichoptic,
eyes separated by broad shiny frons, face narrower than frons; occiput shiny on upper part and faintly pruinescent
on lower part; antenna black, postpedicel long; palpus usually yellowish with darkened apex; labrum about 2X
longer than head height, extensively yellowish, only apex brownish; thorax black, pale setose, 1 npl; postpronotal
lobe and almost entire mesoscutum shiny (except faintly pruinescent narrow marginal edging); legs extensively
yellow, only apical part of femora and tibiae and all tarsi more or less intensively darkened, fore basitarsus slightly
incrassate; wing faintly brownish infuscate, axillary incision very obtuse, anal lobe almost absent, anal vein
incomplete, halter yellow; abdomen shiny, brownish to black dorsally and usually yellowish to brownish yellow
ventrally, pale setose; terminalia directed upwards; cerci separated from each other and from epandrium; cercus with
straight upper margin, apex slightly produced beyond apex of epandrium, covered with setulae; epandrium fused
with hypandrium, with somewhat produced truncate apical part, covered long pale setae, some spinule-like setulae
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at apex; subepandrial sclerite entire, with apex recurved as slender projection, extended beyond lower margin of
epandrium; hypandrium deeply concave medially, with very narrow slightly sclerotised rim anteriorly, produced
as digitiform lobes posterolaterally, lobes covered with long, pale setae; phallus well exposed, pale yellow, mostly
uniformly thin, somewhat thicker on basal part, gently curved; ejaculatory apodeme far before basal curvature of
phallus, very close to base of cerci, with two small lateral wings. Female similar to male, face somewhat broader.
Distribution. Palaearctic. Fennoscandia, mountains of Europe with the exception of southernmost parts,
northwest and north parts of European Russia (Barták 2007; Shamshev 2016).
Rhamphomyia (Pararhamphomyia) armipes Sack
(Figs 28–31)
Rhamphomyia armipes Sack, 1923: 6. Type-locality: Novaya Zemlya (Krestovy Island), Arkhangelskaya Province, Russia.
Other references: Melander, 1928: 382 (catalogue); Frey, 1955a: 469 (revision); Gorodkov & Kovalev, 1969: 624 (key); Chvála
& Wagner, 1989: 297 (catalogue); Shamshev, 2001b: 325 (key); Yang et al., 2007: 168 (catalogue); Shamshev, 2016: 61
(checklist).
Type material examined. LECTOTYPE (here designated in order to fix identity of the species) ♂ (Fig. 28),
labelled (Fig. 29): “Typus [blue label]”; “Novaj. Semlja. Ökland/ Østre Korsä/ (Achaugel bűgt.)/ No 224/ 15
aűg.”; “Rhamphomyia/ armipes Sack/ nov. spec.”; “Gl.1695/ ZM.Oslo”; “pinx [pink label]”; “Typus [pink label]”;
“NHMO: type collection/ 1000327600”; “NHMO/ Norway [data matrix code]”; “LECTOTYPE/ Rhamphomyia/
armipes Sack/ des. Sinclair 2020 [red label]” (NHMO).
PARALECTOTYPE: Berek (Briska) Ösla [?], 18 Aug. No. 235 (1 ♀, NHMO).
Note on types. This species was originally described on the basis of one male and two female specimens.
Diagnosis. This dark, shiny-legged and pale brown setose species is distinguished from other Pararhamphomyia
by the male hind femur and tibia with triangular projections and apex of subepandrial lobe pointed. Female legs
without pennate setae.
Redescription. Wing length 4.5–5.5 mm. Male (Fig. 28). Head with greyish pruinescence on face, frons,
postgena and occiput. Holoptic, eye with ommatidia larger on upper half, smaller on lower half. Frons represented
by very small triangular space below ocellar tubercle and larger subtriangular space above antennae, bare. Face
slightly divergent towards mouthparts; bare, with narrow oral margin shiny. Ocellar triangle dark, pruinescent,
cellar setae broken; 2–3 pairs of postocellar setae slightly shorter and finer than posterior ocellar setae. Occiput with
rows of dark, long, curved postocular setae; postgena with finer, sinuous setae than occiput. Antenna dark brown;
scape and pedicel of equal lengths; postpedicel broken off. Palpus dark, with long, dark setae. Clypeus with greyish
pruinescence; labrum lustrous and dark reddish-brown, 1.5X head height; labellum dark and bearing dark setulae.
Thorax dark in ground-colour, largely densely grey pruinescent; setae long, fine, brownish. Scutum with 2
indistinct, dark grey vittae between acr and dc rows. Pleura clothed in grey pruinescence. Proepisternum with
several long, dark setae on lower section; upper proepisternum in front of spiracle with several long, dark setae;
prosternum bare. Antepronotum with row of short, stout dark setae. Postpronotum with numerous long setae. Scutum
with biserial acr; dc multi-serial, subequal in length to acr, prescutellar setae longer, not differentiated from lateral
setae; numerous undifferentiated presut spal; numerous undifferentiated npl; numerous prealar setae; 4 or more psut
spal similar to prealars; 1 pal with several shorter setae; 4 pairs of sctl. Laterotergite with several long, dark setae.
Anterior and posterior spiracles dark brown with outer black ring.
Legs dark reddish brown, shiny; coxae with faint greyish pruinescence. Coxae with dark setae. All femora
with distinct white ventral pile; fore femur with anteroventral row of setae, with longer posterior setae; mid and
hind femora with 1 anteroventral and 1 posteroventral row of setae, stronger than on fore femur, less than half
corresponding femur width. Hind femur with triangular posteroventral projection beyond mid-length. Fore tibia and
tarsomere lost. Mid tibia with long, erect setae; posterior setae twice as long than anterior setae; with posteroventral
row of strong setae, longer than tibial width. Hind tibia (Fig. 30) with triangular posteroventral projection on basal
third and long posterodorsal setae; 1 long seta in posteroapical comb. Hind tarsomere 1 similar in size to apical
width of hind tibia, dorsal setae shorter than tibial setae; with dense, dark, stout ventral setae.
Wing lightly infuscate with brown veins; all veins complete (except Sc); CuA+CuP at apical fourth extended to
wing margin as faint fold. Pterostigma brownish; basal costa seta absent; anal lobe well-developed. Axillary incision
right angle. Halter dark brown.
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30 · Zootaxa 4848 (1) © 2020 Magnolia Press
Abdomen dark brown with greyish pruinescence; covered with long, pale brown setae laterally, shorter on
tergites dorsally. Tergite 8 similar in length to tergite 7.
Terminalia (Fig. 31, undissected): Epandrium strongly tapered apically; long setae on apical half dorsally and
ventrally. Cercus with broad base, tapered, shorter than subepandrial lobe [right cercus broken at base]. Subepandrial
lobe cylindrical, with pointed apex; clothed in pruinescence. Hypandrium not visible. Phallus with swollen cylindrical
base; lustrous and reddish brown, not forming loops, gently arched between epandrial lamellae [apex not visible].
FIGURES 28–31. Rhamphomyia (Pararhamphomyia) armipes Sack. 28. male habitus, lectotype, lateral view; 29. lectotype
labels; 30. male hind leg, lectotype, anterior view; 31. male terminalia, lectotype, lateral view. Abbreviations: ph—phallus;
sbepand lb—subepandrial lobe.
Female. Similar to male, except basal costa seta present; postoccipital setae stronger; thoracic setae stronger,
more differentiated, with more numerous sctl; mid tibia slightly arched, distinctly shorter than femur, with 1
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anteroventral and 1 posteroventral row of short, strong setae; without pennate setae; CuA+CuP strong to wing
margin.
Remarks. Sack (1923) included illustrations of the male hindleg and male terminalia, the latter upside down. The
basal costal seta is apparently absent in the male lectotype, but is present in the female. Certain features of the female
(presence of basal costal seta; strong CuA+CuP, midleg chaetotaxy) suggest that the female paralectotype is not
conspecific with the male lectotype. Further collections of both sexes of this species are required for confirmation.
The cylindrical subepandrial lobe with pointed apex is similar in form to that of R. hoeli.
Distribution. This species is known only after the type series collected from Novaya Zemlya Archipelago
(Krestovy Island, Serebryanka Fjord, Berkh Island).
Rhamphomyia (Pararhamphomyia) caudata (Zetterstedt)
Rhamphomyza caudata Zetterstedt, 1838: 565. Type locality (by lectotype designation): Kardis, Sweden.
Other references: Frey, 1955a: 472 (revision, lectotype designation); Barták & Danielson, 2007: 107 (type material
examination).
Rhamphomyza aethiops Zetterstedt, 1838: 568. Type locality (by lectotype designation): Lapponia Umensi (probably Lycksele)
(Sweden).
Other references: Barták & Danielson, 2007: 108 (lectotype designation).
Remarks. Rhamphomyia caudata has a somewhat complicated taxonomic history. As a consequence, some
distributional records of this species may be misleading. Rhamphomyia caudata was identified as the single species
of Empididae from Spitsbergen (Boheman 1866; Holmgren 1869; Collin 1923) and was reflected in Empididae
Catalogues and faunal checklists (e.g., Hackman 1968; Chvála & Wagner 1989: 299; Coulson & Refseth 2004: 104;
Yang et al. 2007: 169). We have examined the specimens of R. caudata noted by Holmgren (housed in NHRS) and
found that, actually, they belong to R. longestylata Frey (see below). It should be noted that Collin was somewhat
confused about the synonymy of R. caudata. He believed that R. longestylata Frey was conspecific with R. caudata
(Zetterstedt) (see Collin, 1961: 378). However, they are distinctly two different species (see Frey 1955a: 472; Barták
& Danielson 2008: 107). A note in Collin’s paper on flies collected from Spitsbergen (Collin 1923: 116) indicates
clearly that he in fact had in front of him a male of R. longestylata rather than R. caudata. Our study does not
confirm the presence of R. caudata on the Arctic islands.
Rhamphomyia (Pararhamphomyia) filicauda Henriksen & Lundbeck
(Fig. 32)
Rhamphomyia filicauda Henriksen & Lundbeck, 1917: 608. Type-locality (by lectotype designation): Greenland.
Other references: Sinclair et al., 2019: 36 (lectotype designation, redescription, full list of references).
Rhamphomyia similata Malloch, 1919: 46. Type-locality: Bernard Harbour, Nunavut, Canada.
Material examined. RUSSIA. Chukotka AO (Wrangel Island): Somnitelnaya Bay, on flower of Dryas
integrifolia, spotted tundra, 19.vii.1966, KBG (1 ♀, ZIN); Somnitelnaya Bay, Mineev Mtns, S Sovetskaya Mtn, 400
m, 20.vii.1966, KBG (1 ♀, ZIN); 9 km SE Sovetskaya Mtn, stream shore, 250 m, 12.vii.1972, KBG (1 ♂, ZIN);
environs of Tundrovaya Mtn, 71°18.469′N 179°44.327′W, BT 3, 5–15.vii.2018, U.V. Babiy (1 ♂, ZIN); Perkatkun,
about 30 km N Somnitelnaya Bay, 350 m, on flower of Dryas integrifolia, spotted tundra, 21.vii.1966, KBG (6 ♂,
15 ♀, ZIN); middle flow of Mamontovaya River, 71°10′N 179°45′W, BT 6, 9.vii.2006, OAK (1 ♂, 1 ♀, ZIN); same
locality, BT 11, 8.vii.2006, OAK (1 ♀, ZIN); same locality, BT 13, 30.vi.2015, OAK (1 ♀, ZIN); spurs of Pervaya
Mtn, 71°09′N 179°45′W, BT 12, 28.vi.2015, OAK (1 ♀, ZIN).
Recognition. Mid-sized (wing length 4.8–5.4 mm) blackish brown species. Male (Fig. 32) holoptic; anepisternum
densely pruinescent, scutum densely brownish pruinescent, presutural dc multiserial, long, acr similar but 2–3-
serial; legs robust, dark brown, no modified podomeres, hind femur with short spine-like setae ventrally, all tibiae
with several long bristle-like ad and pd setae, mid basitarsus with 1 long dorsal seta on about middle; wing faintly
infuscate, CuA+CuP incomplete, halter brown; abdomen densely brownish pruinescent, black setose; epandrium
elongate, narrow, constricted in middle, projecting far beyond cercus; cercus V-shaped, strongly constricted in
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32 · Zootaxa 4848 (1) © 2020 Magnolia Press
middle; phallus well-exposed, filamentous, very long, without loops, gently curved beyond apex of epandrium.
Female: anal lobe of wing darker than wing tip, legs without pennate setae.
Distribution. Holarctic; in North America, R. filicauda occurs across the low and high arctic from Greenland
to Yukon (Sinclair et al. 2019); in Eurasia, it is known only from Wrangel Island.
Habitat. A large series of this species was collected only once (in 1966) in the warmer central region of the
island, in dry spotted tundra. In the 2000s, all collections of this species were also limited by dry biotopes of this
part of the island.
Rhamphomyia (Pararhamphomyia) frigida Sinclair, Vajda, Saigusa & Shamshev
(Fig. 33)
Rhamphomyia (Pararhamphomyia) frigida Sinclair et al., 2019: 40. Type locality: Herschel Is., Yukon, Canada.
Material examined. RUSSIA. Chukotka AO (Wrangel Island): middle flow of Neozhidannaya River, 71°01′N,
179°09′E, BT 1, YPT, 12–16.vii.2016, LFV (12 ♂, 18 ♀, ZIN); same locality, BT 2, YPT, 12–16.vii.2016, LFV
(1 ♂, 1 ♀, ZIN); same locality, BT 3, YPT,12–16.vii.2016, LFV (1 ♂, 2 ♀, ZIN); same locality, BT 4, YPT,
12–16.vii.2016, LFV (2 ♂, ZIN); same locality, BT A, YPT, 8–9.vii.2019, OAK (1 ♂, ZIN); same locality, BT В,
Sw, 8.vii.2019, OAK (3 ♂, 1 ♀, ZIN); same locality and biotope, YPT, 8–9.vii.2019, OAK (2 ♀, ZIN); basin of
Khischniki River, 7 km SE Sovetskaya Mtn, 15.vii.1972, KBG (1 ♂, ZIN). USA. Alaska: Inaru River, 28.vii.1956;
ex. dandelion flowers (1 ♀, CNC).
Recognition. Mid-sized (wing length 4.2–5 mm), dark-legged, greyish species. Male (Fig. 33) holoptic; scutum
densely greyish pruinescent, with two indistinct darker vittae between rows of acr and dc (dorsal view); thorax black
setose, upper proepisternum in front of spiracle bare; main mesonotal setae well differentiated, strong, notopleuron
with cluster of several long blunt-tipped setae, acr biserial, presutural dc 1–2-serial, 6 sctl; spiracles brown; fore
coxa with strong blunt-tipped setae anteriorly; hind tibia evenly thickened toward apex, hind basitarsus slightly
thickened, both with similar long, dense fine setae dorsally; wing faintly infuscate, CuA+CuP incomplete, basal
costal seta absent, halter with yellowish knob; abdomen faintly greyish pruinescent, with yellowish setae; terminalia
with subepandrial lobe, phallus mostly exposed, thickened at base, without loops, evenly curved. Female similar
to male, except eyes dichoptic, wing darker infuscate, legs and abdomen with much shorter setation; legs without
pennate setae, hind tibia and basitarsus slender.
Distribution. Holarctic; in North America this species is known from a few localities along the Beaufort Sea and
the Pribilof Islands in the Bering Sea (Sinclair et al. 2019); in Eurasia, it is recorded only from Wrangel Island.
Habitat. Almost all specimens of this species were collected in the 2000s in the colder region of Wrangel Island
(Neozhidannaya River), in biotopes with different moisture.
Rhamphomyia (Pararhamphomyia) hoeli Frey
(Fig. 34)
Rhamphomyia (Pararhamphomyia) hoelsi Frey, 1950: 100. Type-locality: Greenland.
Rhamphomyia hoeli: Frey, 1955b: 482 (revision); Downes, 1970: 779 (behaviour); Danks, 1981: 465 (arctic insects); Yang
et al., 2007: 171, 195 (catalogue); Barták, 2015: 576 (Greenland fauna); Sinclair et al., 2019: 50 (lectotype designation,
redescription).
Material examined. Russia. Chukotka AO (Wrangel Island): Somnitelnaya Bay, on flower of Dryas integrifolia,
spotted tundra, 19.vii.1966, KBG (2 ♂, 2 ♀, ZIN); environs of pass Vjuchny, NW Somnitelnaya Bay, 21.vii.1972,
KBG (1 ♂, ZIN); 5 km N Somnitelnaya Bay, valley of Somnitelnaya River, small meadow, 25.vii.1972, KBG (2 ♀,
ZIN); same locality, 7–8.vii.1972, KBG (2 ♀, ZIN); Somnitelnaya Bay, valley of Somnitelnaya River, Mineev Mtns,
150 m, pebbles, on flower of Potentilla emarginata, 22.vii.1966, KBG (7 ♂, 3 ♀, ZIN); near north slope of Berri Peak,
6 km ESE Sovetskaya Mtn, 15.vii.1972, KBG (5 ♂, 2 ♀, ZIN); upper flow of Khischniki River, 7 km SE mountain
Sovetskaya, Dryas tundra near river, 200 m, 11.vii.1972, KBG (7 ♂, 2 ♀, ZIN); middle flow of Mamontovaya River,
N Perkatkun, osier-bed in river valley, 17,22.vii.1972, KBG (8 ♂, 1 ♀, ZIN); same locality, 71°10′N 179°45′W, BT
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 33
20A, YPT, 2–4.vii.2015, OAK (1 ♂, ZIN); spurs of Pervaya Mtn, 71°09′N, 179°27′W, BT 11, Sw, 28.vi.2015, OAK
(2 ♂, ZIN); same locality, BT 14, Sw, 28.vi.2015, OAK (1 ♂, ZIN); upper flow of Neizvestnaya River, 71°13′N,
179°19′W, BT 5А, 26.vi.2006, OAK (1 ♂, ZIN); same locality and biotope, 26.vi.2015, OAK (1♂, ZIN).
FIGURES 32–34. Rhamphomyia (Pararhamphomyia), male habitus, lateral view. 32. R. (P.) filicauda Henriksen & Lundbeck;
33. R. (P.) frigida Sinclair, Vajda, Saigusa & Shamshev; 34. R. (P.) hoeli Frey.
Recognition. Rather small species (wing length 3.5–4.2 mm) with dark legs and densely pruinescent thorax
and abdomen. Male (Fig. 34) holoptic; antenna dark brown; scutum brownish grey with two indistinct darker vittae
between rows of acr and dc (dorsal view); thorax dark setose, upper proepisternum in front of spiracle with 0–2
short setae; mesonotal setae well differentiated, bristle-like, acrostichals and dorsocentrals long, 1–2-serial, 2–3 npl,
4 sctl; spiracles dark brown; legs dark setose, fore coxa with fine setae anteriorly, tibiae and tarsomeres slender;
mid tibia with 2–3 anterodorsal setae (except 1 long seta of subapical circlet), about 2X as long as tibial width; hind
tibia with moderately long strong setae dorsally, 1 seta in posteroapical comb; hind femur with complete row of
anteroventral setae (longest subapical setae nearly as long as femur width), hind basitarsus only with short setae
dorsally; wing whitish, CuA+CuP evanescent apically, not quite reaching wing-margin or extended as faint fold,
basal costa seta present, axillary incision right-angled, halter pale brown to yellow; abdomen greyish, paler than
thoracic pleuron, dark setose; terminalia small; cercus shorter than epandrium, with broad base, tapered at mid-
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length to narrow apical portion, covered with short setulae; subepandrial lobe present, cylindrical, with pointed
apex, projecting slightly farther than cercus; epandrium subtriangular, tapered to truncate, slightly bilobed setose
apex, curved medially, outer apex with cluster of long, dark stout setae; hypandrium short, boot-shaped, ending
in middle of swollen phallus base; phallus short, gently arched between epandrial lamellae, not forming loops,
base swollen, apex with shallow S-shaped curvature. Female similar to male, except eyes dichoptic; wing slightly
infuscate, CuA+CuP entirely sclerotized, brownish; leg chaetotaxy reduced, without pennate setae.
Distribution. Holarctic; in North America, R. hoeli is known from the low and high arctic region east of
Hudson Bay and Greenland (Sinclair et al. 2019); in Eurasia, it is recorded only from Wrangel Island.
Habitat. On Wrangel Island this species was collected mainly in valleys of rivers and streams as well as some
wet biotopes.
Rhamphomyia (Pararhamphomyia) kjellmanii Holmgren
(Fig. 35)
Rhamphomyia kjellmanii Holmgren, 1880: 22. Type locality (by lectotype designation): Novaya Zemlya, Arkhangelskaya
Province, Russia.
Other references: Holmgren & Aurivillius, 1883: 163 (expedition results); Bezzi, 1903: 226 (catalogue); Melander, 1928: 195
(catalogue).
Rhamphomyia kjellmani Holmgren: Frey, 1915: 3, 8 (faunistic survey).
Rhamphomyia (Pararhamphomyia) kjellmanii Holmgren: Frey, 1955b: 500 (revision); Chvála & Wagner, 1989: 301 (catalogue);
Yang et al., 2007: 172 (catalogue); Shamshev, 2016: 64 (checklist); Shamshev & Sinclair, 2018: 311 (lectotype designation,
redescription); Sinclair et al., 2019: 54 (revision).
Rhamphomyia uralensis Becker, 1915: 58. Type-locality (by lectotype designation): mouth of Kara River (about 69°02′N
64°35′E), Yamalo-Nenets, Russia.
Rhamphomyia (Pararhamphomyia) uralensis Becker: Frey, 1955b: 500 (revision); Chvála & Wagner, 1989: 306 (catalogue); Yang
et al., 2007: 178 (catalogue); Shamshev, 2016: 71 (checklist); Sinclair et al., 2019: 54 (synonym, lectotype designation).
Material examined. RUSSIA. Archangelskaya Prov. (Novaya Zemlya Archipelago): steamer deck, in sea near
western shore of Novaya Zemlya, 20.vii.1948, Korotkevich (1 ♂, 1 ♀, ZIN). Chukotka AO (Wrangel Island):
middle flow of Mamontovaya River, 71°10′N 179°45′W, BT 20A, Sw, 2.vii.2015, OAK (1 ♂, ZIN); same locality
and biotope, YPT, 2–4.vii.2015, OAK (3 ♂, ZIN); same locality, BT 3, Sw, 1.vii.2015, OAK (3 ♂, ZIN); same
locality, BT 2, YPT, 2–4.vii.2015, OAK (1 ♂, ZIN); same locality, BT 1A, Sw, 2.vii.2015, OAK (1 ♂, ZIN);
same locality and biotope, YPT, 2–4.vii.2015, OAK (1 ♂, 2 ♀, ZIN); same locality, BT 7, Sw, 1.vii.2015, OAK (1
♂, ZIN); upper flow of Neizvestnaya River, 71°13′N, 179°19′W, BT 12, Sw, 6.vii.2015, OAK (7 ♂, ZIN); same
locality, BT 13, YPT, 6–15.vii.2015, OAK (1 ♂, ZIN); same locality, BT 7A, PT, 25.vi.–10.vii.2015, OAK (1 ♂,
ZIN); same locality, BT C1, Sw, 24.vi.2015, OAK (1 ♂, ZIN). Chukotka AO (Kolyuchin Island): 25–27.vii.1938,
Hecker (4 ♂, 8 ♀, ZIN).
Recognition. Mid-sized (wing length 4.2–4.5 mm) species. Male (Fig. 35) holoptic; scutum densely brownish
grey pruinescent, dc uniserial, acr biserial, notopleuron with 4–6 long strong blunt-tipped setae, 6 sctl, laterotergite
with pale setae; legs dark brown, shiny, fore coxa with strong blunt-tipped setae anteriorly, hind tibia clavate and
clothed in dense long hair-like setae, hind tarsomeres 1–2 thickened, with similar setae dorsally; wing faintly
infuscate, CuA+CuP incomplete, halter yellowish (sometimes dirty yellow); abdomen faintly greyish pruinescent,
covered with pale setae; terminalia: cercus short, one-third length of epandrium, tapered apically; subepandrial
lobe present, longer than cercus, clavate; epandrium gradually tapered, bearing numerous long, pale setae apically;
phallus well exposed, slender, gently sinuate on about middle, looping beyond epandrium and curving back to
cercus. Female wing faintly brownish infuscate, hind tibia and tarsus slender, legs with simple short setae.
Distribution. Holarctic; in Eurasia, R. kjellmanii extends across arctic and subarctic Russia, including both
continental areas (Taymyr, Yakutia, Chukotka) and islands (Novaya Zemlya, Wrangel, Kolyuchin). In North
America, this species is known only from the extreme northwestern region (Sinclair et al. 2019).
Habitat. On Wrangel Island this species was collected mainly in wet and, less commonly, in moderately
moistened habitats in the warmest central part of the island.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 35
FIGURE 35. Rhamphomyia (Pararhamphomyia) kjellmanii Holmgren, male habitus, lateral view.
Rhamphomyia (Pararhamphomyia) longestylata Frey
(Figs 36, 37)
Rhamphomyia longestylata Frey in Poppius et al., 1917: 682. Type localities: “Schwedisch-Lappland, Sarek: Pårte, Kåtokjokk”,
Sweden.
Rhamphomyia longestylata Frey in Lundström & Frey, 1913: 8 (nomen nudum).
Rhamphomyia (Pararhamphomyia) longestylata Frey: Frey, 1922: 44 (revision); Melander, 1928: 196 (catalogue); Frey,
1955b: 485 (revision); Jonassen et al., 2013: 214 (faunistic survey); Kahanpää, 2014: 187 (checklist); Shamshev, 2016: 65
(checklist).
Rhamphomyia caudata: authors, nec Zetterstedt, 1838: Boheman, 1866: 570 (Rhamphomyza); Holmgren, 1869: 26; Collin,
1923: 116 (expedition results); Collin, 1961: 378 (revision); Hackman, 1968: 93 (faunistic survey); Chvála & Wagner,
1989: 299 (catalogue); Coulson & Refseth 2004: 104 (checklist Svalbard); Yang et al., 2007: 169 (catalogue).
Material examined. NORWAY. Spitsbergen (Svalbard Archipelago): Advent Bay, Holmgren (5 ♂, 6 ♀, NHRS);
Mimerdalen, Pyramiden, tundra, 29.viii.1979, KBG; on flowers of Dryas octopetala (1 ♂, 1 ♀, ZIN).
Diagnosis. Mid-sized (wing length about 4 mm), blackish, shiny flies. Male (Fig. 36) holoptic; thorax and
abdomen black setose, scutum and anepisternum shiny; legs dark brown, hind basitarsus slightly uniformly
thickened, all tibiae and tarsi clothed in long numerous fine setae but hind tibia with long setae on about apical half
only; abdomen shiny, epandrium and cerci elongate, cercus finger-like beyond dorsal cavity; phallus well exposed,
filamentous, long, gently curved. Female with brownish infuscate wing, short setose slender legs without pennate
setae.
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36 · Zootaxa 4848 (1) © 2020 Magnolia Press
Redescription. Wing length 3.7–4.1 mm. Male (Fig. 36). Head with faint greyish pruinescence on face, frons,
postgena, ocellar triangle and occiput; dark setose. Holoptic; upper ommatidia enlarged. Frons represented by very
small subtriangular space below ocellar tubercle and larger subtriangular space above antennae, bare. Face broad,
bare. Ocellar triangle with 2 long fine and some short setae. Occiput with numerous long fine setae, including
postoculars; postgena with numerous hair-like setae. Antenna blackish; scape short, slightly longer than pedicel,
both with short setulae; postpedicel conical, with slightly concave ventral margin on apical part, nearly 2.5X longer
than wide; stylus short, slightly shorter than postpedicel basal width. Palpus dark; with several long, dark fine setae.
Proboscis with labrum dark reddish-brown, nearly 1.3X as long as head height.
Thorax dark in ground-colour, with black setation; prothoracic sclerites mostly densely greyish pruinescent,
antepronotum and postpronotal lobe faintly pruinescent, subshiny; mesonotum almost entirely shiny, only notopleuron
along lower margin, postalar tubercle, scutum in front of scutellum and scutellum very faintly pruinescent; mesopleuron
mostly greyish pruinescent, anepisternum shiny, also usually katepisternum on upper part and anepimeron anteriorly
shiny. Proepisternum with few moderately long fine setae on lower section; upper proepisternum in front of spiracle
without setae. Prosternum bare. Antepronotum with several short setae. Postpronotal lobe with 1–2 long, stronger
and several shorter fine setae. Mesonotal setation: presutural supra-alar space with several long fine setae, 3–4 npl
(with numerous finer setae anteriorly), 1–2 psut spal (with additional setulae anteriorly), 2 pal (often with additional
setulae), 4–6 sctl (usually 6, variable in length and robustness, sometimes asymmetrically set); acr short, fine,
arranged in 2 close irregular rows, lacking on prescutellar depression; presutural dc longer than acr, fine, arranged
in 2 irregular rows, postsutural dc 1–2-serial, 3–4 prescutellars longest. Laterotergite with several long fine setae.
Anterior and posterior spiracles brown.
Legs slender, uniformly brownish; mostly shiny, coxae and trochanters faintly greyish pruinescence; black
setose. Fore coxa covered with long fine setae anteriorly. All tibia slightly evenly thickened toward apex; hind
basitarsus slightly uniformly thickened, nearly as broad as hind tibia width at apex. All femora whitish pilose
ventrally, with very short setation; fore and mid femora with anteroventral and posteroventral rows of short fine
setae; hind femur with row of short anteroventral setae. Fore and mid tibiae with numerous long fine setae dorsally
and posteriorly (about 2X tibial width); mid tibia with numerous stronger long anteroventral and posteroventral
setae on about apical 2/3; hind tibia with long fine setae dorsally on about apical half, 1 long seta in posteroapical
comb. Tarsomeres 1–2 of all legs with numerous long setae dorsally (somewhat longer on basitarsi).
Wing membrane faintly infuscate, with brownish veins; CuA+CuP (anal vein) incomplete, evanescent apically.
Pterostigma brownish yellow. Basal costa seta absent. Anal lobe well-developed; axillary incision right-angled or
slightly obtuse. Squama brown, dark fringed. Halter brown.
Abdomen dark brown, shiny; mostly with black fine moderately long setae. Segments 6–7 unmodified. Segment
8 with almost fused tergite and sternite; tergite 8 simple, entire, about half length of sternite 7, with straight posterior
margin, with numerous long setae along posterior margin; tergite 8 simple, enlarged, subrectangular in lateral view,
V-shaped viewed posteriorly, covered with numerous long setae posteriorly.
Terminalia (Fig. 37) dark brown, directed oblique upstairs. Cerci separated from each other and from epandrium,
not extended beyond tergite 8 anteriorly; cercus long, narrow, broadly concave on upper margin, apical portion finger
like, parallel-sided (lateral view), at least as broad as middle portion, with small internal subtriangular projection
slightly beyond middle (dorsal view); covered with short fine setae; without pair of subepandrial lobes beneath.
Epandrium subrectangular, elongate, about 4X as long as broad, broadly concave on upper margin, with straight
lower margin, somewhat broadened and angularly expanded apically, extended slightly beyond cerci posteriorly;
with cluster of numerous long dark setae near upper anterior corner, covered with dense fine setulae on broadened
apical part, some numerous moderately long setae along lower margin. Hypandrium very narrow, rim-like, entire,
well sclerotized; bare. Phallus well exposed, long; mostly hair-like, knob-like thickened just beyond hypandrium;
with right-angle curvature shortly beyond basal thickening, otherwise gently arcuate. Ejaculatory apodeme not
extended beyond basal curvature of phallus, with broad lateral wings and narrower vertical wing.
Female. Similar to male, except dichoptic, frons broad, shiny, with marginal setulae; wing brownish uniformly
infuscate; leg with tibiae and tarsi slender, with short setation, without pennate setae; abdomen covered with short
setae; cercus long slender, with dark setulae.
Distribution. Palaearctic; R. longestylata is recorded only from northern Scandinavia (Finland, Norway,
Sweden), nearby regions of Russia (Murmanskaya Province, Nenets) and Spitsbergen Island (Norway).
Remarks. Rhamphomyia longestylata is assigned to the R. caudata group (Sinclair et al. 2019) or to the R.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 37
longestylata group (Barták & Kubík 2009). It is very similar to R. ursinella and the main differences between these
species are provided in the key. We have examined the syntypes of R. longestylata (housed in MZH and NHRS), but
the lectotype has not been designated.
FIGURES 36–38. Rhamphomyia (Pararhamphomyia). 36. R. (P.) longestylata Frey, male habitus, lateral view; 37. same
species, male terminalia, lateral view; 38. R. (P.) nordqvistii Holmgren, female habitus, lectotype, lateral view (from Shamshev
& Sinclair 2018: 314, fig. 10). Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium; hypd—
hypandrium; ph—phallus.
Rhamphomyia (Pararhamphomyia) nordqvistii Holmgren
(Fig. 38)
Rhamphomyia nordqvistii Holmgren, 1880: 23. Type locality (by lectotype designation): Novaya Zemlya, Arkhangelskaya
Province, Russia.
Other references: Holmgren & Aurivillius, 1883: 164 (expedition results).
Rhamphomyia nordquisti Holmgren: Frey, 1915: 3, 8 (faunistic survey); Sack, 1923: 7 (expedition results).
Rhamphomyia nordquistii Holmgren: Bezzi, 1903: 228 (catalogue); Melander, 1928: 200 (catalogue).
Rhamphomyia (Pararhamphomyia) nordqvisti Holmgren: Chvála & Wagner, 1989: 303 (catalogue); Shamshev, 2001b: 325
(key); Yang et al., 2007: 174 (catalogue).
Rhamphomyia (Pararhamphomyia) nordqvistii Holmgren: Shamshev, 2016: 66 (checklist); Shamshev & Sinclair, 2018: 313
(lectotype designation, redescription).
Rhamphomyia (Pararhamphomyia) eunordquisti Frey, 1922: 41 (as var. of R. obscura Zetterstedt). Type localities: “von Nord-
SHAMSHEV ET AL.
38 · Zootaxa 4848 (1) © 2020 Magnolia Press
Sibirien (Dudinka) und Kamtschatka”, Russia.
Other references: Melander, 1928: 201 (catalogue); Frey, 1955b: 490 (revision); Chvála & Wagner, 1989: 303 (catalogue); Yang
et al., 2007: 174 (catalogue).
Rhamphomyia (Pararhamphomyia) eunordqvisti Frey: Frey, 1935: 2 (expedition results).
Recognition. This species remains known by the female sex only. Small (wing length of lectotype 3.2 mm) blackish
flies (Fig. 38); thorax black setose, upper proepisternum in front of spiracle with 3 short setae; scutum brownish
grey pruinescent; main mesonotal setae fine, hardly distinguishable, 4 npl, 6 sctl (4 stronger), presutural dc rather
long, 2–3-serial, acr biserial; legs brownish, with simple setae, fore coxa with fine setae anteriorly; wing uniformly
brownish infuscate, no distinct basal costal seta, CuA+CuP incomplete, axillary incision right-angled, halter
brownish; abdomen dark brown, faintly greyish pruinescent, subshiny (lateral view), with brownish setae (extracted
from Shamshev & Sinclair 2018: 313).
Distribution. This species remains known only from Novaya Zemlya. In addition to the type material, Sack
(1923: 7) noted a female of R. nordqvistii among his specimens of Empididae collected from Krestovy Island
(Novaya Zemlya). However, this record needs verification because the original description of R. nordqvistii is non-
informative and Sack never examined the types of this species. Also, Frey (1915: 8) identified as R. nordqvistii
several females from the delta of Lena River (Yakutia). At least one specimen of this series is present in ZIN and it
belongs to a different species.
Remarks. Shamshev & Sinclair (2018) have already noted that the synonymy of R. eunordquisti with R.
nordqvistii is doubtful.
Rhamphomyia (Pararhamphomyia) septentrionalis Sinclair, Vajda, Saigusa & Shamshev
(Figs 39, 40)
Rhamphomyia (Pararhamphomyia) septentrionalis Sinclair et al., 2019: 65. Type locality: Victoria Is. (71°17′N, 114°W),
Northwest Territories, Canada.
Other references: Shamshev, 2016: 70 (checklist, partly as Rhamphomyia (Pararhamphomyia) tenuiterfilata Becker, 1900).
Material examined. RUSSIA. Chukotka AO (Wrangel Island): middle flow of Mamontovaya River, N
Perkatkun, osier-bed in river valley, 17, 21.vii.1972, KBG (17 ♂, 13 ♀, ZIN); same locality, 71°10′N 179°45′W, BT
14, 9.vii.2006, OAK (4 ex., SZM); same locality and biotope, Sw, 2.vii.2015, OAK (2 ♂, 5 ♀, ZIN); same locality,
BT 3, 9.vii.2006, OAK (1 ex., SZM); same locality and biotope, Sw, 1.vii.2015, OAK (1 ♂, ZIN); same locality,
BT 20A, Sw, 2.vii.2015, OAK (2 ♂, ZIN); spurs of Pervaya Mtn, 71°09′N, 179°27′W, BT 11, Sw, 28.vi.2015, OAK
(6 ♂, ZIN); upper flow of Neizvestnaya River, 71°13′N, 179°19′W, BT 4A, 5.vii.2006, OAK (1 ex., SZM); same
locality and biotope, Sw, 22.vi.2015, OAK (3 ♂, ZIN); same locality and biotope, 26.vi.2015, OAK (1 ♂, ZIN).
Recognition. Mid-sized (wing length 5.4–6.1) blackish slender flies. Male (Fig. 39) holoptic; scutum faintly
greyish pruinescent, with 4 narrow shiny vittae; presutural dc pale 1–2-serial, hair-like, moderately long, acr biserial,
similar to but somewhat shorter than dc, laterotergite with pale setae; legs long, very slender, shiny, with strongly
reduced setation; hind tibia gently curved inwards beyond middle, hind basitarsus long spindle-shaped, with dense
long hair-like setae dorsally; wing faintly infuscate, CuA+CuP almost complete, well sclerotized, disappearing or
evanescent just before wing margin, halter yellow; abdomen with shiny dorsum, covered with pale setae; terminalia
longer than thorax; epandrium elongate, almost strap-like, somewhat broadly constricted on about middle, with
broadly rounded apex; cercus long, but shorter than epandrium, narrow, constricted on about middle; phallus very
long, hair-like, without loops, extended beyond epandrium. Female (Fig. 40) wing broad, brownish; hind tibia and
tarsus simple; legs without pennate setae.
Distribution. Holarctic; R. septentrionalis occurs in the low arctic west of Hudson Bay of North America and
in northern regions of Russia (Sinclair et al. 2019).
Habitat. On Wrangel Island this species was collected in the central part of the island, where it was found
mainly in river valleys with willow bushes.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 39
FIGURES 39, 40. Rhamphomyia (Pararhamphomyia) septentrionalis Sinclair, Vajda, Saigusa & Shamshev, habitus, lateral
view. 39. male; 40. female.
Rhamphomyia (Pararhamphomyia) subfilicauda Shamshev & Sinclair sp. nov.
(Figs 41, 42)
urn:lsid:zoobank.org:act:27AE8916-35F9-42A2-BD16-692DADF550F9
Type material. HOLOTYPE ♂, labelled: [RUSSIA, Chukotka AO (Wrangel Island)] “upper flow of river
Neizvestnaya, 71°13′N 179°19′W, BT 2, 3.vii.2006, O.A. Khruleva”; “Holotypus/ Rhamphomyia/ subfilicauda/
Shamshev, Sinclair sp. n. [red label]” (INS_DIP_0000621, ZIN; terminalia dissected, in microvial pinned with
specimen).
Diagnosis. Mid-sized (wing length 4.3 mm) blackish flies of the R. pusilla group. Scutum uniformly velvety
brown, without vittae; postpronotal lobe and mesonotum with numerous uniformly very long fine setae (including
acr and dc), main mesonotal setae undifferentiated; hind tibia curved inwards closer to base; halter brownish; wing
whitish, CuA+CuP complete; phallus well exposed, very long, mostly hair-like.
Description. Male (Fig. 41). Wing 4.3 mm, body 4 mm. Head with brownish pruinescence on face, frons,
postgena, ocellar triangle and occiput; dark setose. Eyes holoptic, touching on frons. Ocellar triangle with several
long fine setae. Occiput with numerous uniformly long thin setae, including postoculars. Antenna blackish brown;
scape and pedicel short, scape slightly longer, both with short setulae; postpedicel conical, 3X longer than wide;
stylus very short, slightly shorter than postpedicel basal width. Palpus dark; with numerous long, dark fine setae.
Labrum 1.3X head height.
Thorax densely brownish pruinescent; with black setation; scutum uniformly velvety brown pruinescent, without
vittae. Proepisternum with tuft of several long fine setae on lower section; upper proepisternum in front of spiracle
without setae. Prosternum bare. Antepronotum with numerous short setae. Postpronotal lobe and mesonotum with
numerous uniformly very long fine setae (including acr and dc), main mesonotal setae undifferentiated; 6 sctl;
acr arranged in 2 irregular rows, absent on prescutellar depression; presutural dc separated by bare space from
supra-alars, arranged in 3–4 irregular rows, postsutural dc 1–2-serial. Laterotergite with numerous long fine setae.
Anterior and posterior spiracles brown.
Legs rather robust; mid legs and hind tarsi missing, remaining parts uniformly dark brown, black setose. Fore
basitarsus slightly expanded; hind tibia straight, evenly expanded towards apex. Fore femur with rows of moderately
long, fine anteroventral (becoming longer toward base of femur) and mostly minute (3–4 moderately long setae near
extreme base) posteroventral setae, some moderately long setae posteriorly; hind femur with moderately long to
long (on about apical half of femur) setae anteroventrally, some moderately long setae anteriorly. Fore tibia with
SHAMSHEV ET AL.
40 · Zootaxa 4848 (1) © 2020 Magnolia Press
numerous long setae dorsally and posterodorsally (longest setae nearly 1.5X longer than tibia width); fore basitarsus
with similar setae; hind tibia with numerous very long setae dorsally and similar setae anteroventrally (except
extreme apex and base), 1 seta in posteroapical comb.
Wing membrane whitish; CuA+CuP (anal vein) incomplete; 1 basal costal seta present; axillary incision almost
90°. Squama brownish, dark fringed. Halter brownish.
Abdomen densely brownish grey pruinescent, covered with long black setae (shorter on tergites dorsally).
Segments 6–7 with unmodified structure; tergite 6 covered with only scattered minute setulae; tergite 7 mostly
with minute setulae, bearing several fine short setae posteromarginally and laterally. Segment 8 with tergite and
sternite separated; tergite 8 simple, entire, subtriangular viewed laterally, very narrow mid-dorsally, broadly concave
anteriorly, with almost straight posterior margin, numerous very long setae along posterior margin; sternite 8 simple,
enlarged, subrectangular in lateral view, V-shaped viewed posteriorly, covered with numerous very long setae.
Terminalia (Fig. 42) dark brown, black setose. Cerci separated from each other and from epandrium, not extended
beyond tergite 8 anteriorly; cercus rather short, deeply cleft mid-dorsally and divided almost entirely into two parts:
basal part subrectangular (lateral view), apical part rather elongate oval, both covered with short fine setae; without
pair of subepandrial lobes beneath. Epandrium subtriangular, with strongly produced narrow apical part, slightly
concave on upper margin, with more or less straight lower margin, extended far beyond cerci posteriorly; with
row of 6 long fine closely set setae near upper anterior corner, cluster of short, very closely set spine-like setae on
upper part slightly beyond middle, numerous long setae over lower margin becoming denser and stronger slightly
beyond cluster of spine-like setae but short and scattered on remaining part of epandrium. Subepandrial sclerite
produced slightly beyond lower margin of epandrium. Hypandrium very narrow, rim-like, entire, well sclerotized;
bare; gonocoxal apodeme directed anteriorly. Phallus well exposed, very long; mostly hair-like, knob-like thickened
just beyond hypandrium; slightly sinuate and with broad even curvature shortly beyond basal expansion, gently
sinuate on remaining part. Ejaculatory apodeme not extended beyond basal curvature of phallus, with broad lateral
wings and somewhat narrower vertical wing.
Female. Unknown.
Distribution. Eurasia; only Wrangel Island.
Etymology. The specific name refers to the similarity of the new species to R. filicauda Henriksen &
Lundbeck.
FIGURES 41, 42. Rhamphomyia (Pararhamphomyia) subfilicauda sp. nov., male, holotype. 41. habitus, lateral view; 42.
terminalia, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium; hypd—hypandrium;
ph—phallus.
Remarks. The new species belongs to the R. pusilla group sensu Barták & Kubík (2009) (see also Sinclair et
al. 2019: 65). It is similar to R. filicauda Henriksen & Lundbeck, the main differences between these species have
been provided in the key. In addition, the new species could be compared with R. aversa Frey known only from
Tajikistan. However, R. aversa differs from the new species primarily by faintly infuscate wings (vs. whitish).
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 41
Habitat. The single specimen was collected in the moss-herb-dryad tundra in the warmer region of Wrangel
Island.
Rhamphomyia (Pararhamphomyia) submacrura Shamshev & Sinclair sp. nov.
(Figs 43, 44)
urn:lsid:zoobank.org:act:6D5C99D5-DC42-4972-857E-D9A31B87234A
Type material. HOLOTYPE ♂, labelled: [RUSSIA: Chukotka AO (Wrangel Island)] “Somnitelnaya Bay,/
south o. [= ostrov, island] Wrangel/ Gorodkov 20.vii.[1]966”; “400 m/ Mineev Mountains, S g. [= gora, mountain]
Sovetskaya”; “Holotypus/ Rhamphomyia/ submacrura/ Shamshev, Sinclair sp. n. [red label]” (INS_DIP_0000615,
ZIN; terminalia dissected, in microvial pinned with specimen).
Diagnosis. Rather large brownish grey species of the R. pusilla group with long, slender legs, curiously long
epandrium and, especially, phallus; presutural dc setae multiserial, halter brownish; hind tibia and basitarsus only
slightly thickened; phallus extremely long, hair-like.
Description. Male (Fig. 43). Wing length 6.7 mm, body length about 6.5 mm.
Head with brownish grey pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Eyes
holoptic, touching on frons. Ocellar triangle with several long fine setae. Occiput with numerous uniformly long
thin setae, including postoculars. Antenna blackish brown; scape and pedicel short, scape slightly longer, both with
short setulae; postpedicel conical, 3X longer than wide; stylus short, slightly shorter than postpedicel basal width.
Palpus dark; with numerous long, dark fine setae. Labrum nearly of head height.
Thorax densely brownish grey pruinescent; with black setation; scutum uniformly brownish pruinescent,
without vittae. Proepisternum with tuft of several long fine setae on lower section; upper proepisternum in front
of spiracle without setae. Prosternum bare. Antepronotum with numerous fine setae shorter dorsally and longer
laterally. Postpronotal seta hardly distinguishable; postpronotal lobe with numerous setae of different lengths and
robustness (1–2 long setae slightly stronger). Mesonotal setation: presutural supra-alar space covered with fine
mostly short setae (1–2 setae longer), 4 npl (with numerous shorter and finer setae anteriorly), 1 sut spal (with
several shorter setae anteriorly), 2 pal, 14 sctl; acr short fine, arranged in 2–3 irregular rows, lacking on prescutellar
depression; presutural dc longer than acr, multiserial, not separated by bare space from supra-alars, postsutural dc
arranged in 2–3 irregular rows, several prescutellars longest. Laterotergite with numerous long fine setae. Anterior
and posterior spiracles brown.
Legs long, slender, uniformly brownish, subshiny, black setose. Hind tibia somewhat evenly thickened toward
apex; hind basitarsus slightly thickened but somewhat narrower that hind tibia at apex, cylindrical, 1.7X shorter
than hind tibia. Fore coxa with numerous long simple setae anteriorly. Femora whitish pilose ventrally. Fore femur
with rows of fine moderately long anteroventral and posteroventral setae. Mid femur with numerous short strong
setae ventrally. Hind femur with row of anteroventral setae, row with short, fine setae on basal half but longer and
stronger setae on apical half; spinule-like ventral setae on basal part. Fore tibia with numerous long, quite strong
setae posterodorsally and posteriorly (longest setae nearly 1.5X tibia width). Mid tibia with several moderately
long anteroventral and posteroventral setae, some strong spine-like setae toward apex ventrally. Hind tibia with
numerous moderately long, rather strong setae over entire length; 1 seta in posteroapical comb. Fore basitarsus with
some fine moderately long setae posterodorsally, remaining fore tarsomeres, mid and hind tarsi covered with simple
short setae.
Wing membrane faintly infuscate; CuA+CuP (anal vein) complete, well sclerotised; basal costal seta absent;
axillary incision acute. Squama brownish, dark fringed. Halter brownish.
Abdomen dark brown, brownish grey pruinescent, tergites somewhat lustrous; covered with long, dense, fine
black setae. Segments 6–7 with unmodified structure; tergite 6 only with scattered minute setulae dorsally and
several short fine posteromarginal setae; tergite 7 almost bare, with some fine setae posteromarginally (longer mid-
dorsally) and laterally. Segment 8 with tergite and sternite separated; tergite 8 simple, entire, subtriangular viewed
laterally, very narrow mid-dorsally, broadly concave anteriorly, with almost straight posterior margin, numerous
very long setae along posterior margin; sternite 8 simple, enlarged, subrectangular in lateral view, V-shaped viewed
posteriorly, covered with numerous very long setae.
SHAMSHEV ET AL.
42 · Zootaxa 4848 (1) © 2020 Magnolia Press
FIGURES 43, 44. Rhamphomyia (Pararhamphomyia) submacrura sp. nov., male, holotype. 43. habitus, lateral view; 44.
terminalia, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium; hypd—hypandrium;
ph—phallus.
Terminalia (Fig. 44) dark brown, black setose. Cerci separated from each other and from epandrium, not
extended beyond tergite 8 anteriorly, without pair of subepandrial lobes beneath; cercus rather short, deeply cleft mid-
dorsally and divided almost entirely into two parts; basal part subquadrate (lateral view); apical part subrectangular,
elongate, narrower on basal section, somewhat broadened apically, apex slightly concave; both parts covered with
short fine setae. Epandrium subtriangular, with strongly produced narrow apical part, nearly 4X as long as basal
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 43
portion width, extended far beyond cerci posteriorly, slightly concave on upper margin, with more or less straight
lower margin; with several long closely set setae near upper anterior corner, numerous long setae over lower margin
becoming denser and stronger on about middle, moderately long and scattered setae on apical part of epandrium.
Subepandrial sclerite produced slightly beyond lower margin of epandrium. Hypandrium very narrow, rim-like,
entire, well sclerotized; bare; gonocoxal apodeme directed anteriorly. Phallus well exposed, extremely long; mostly
hair-like, knob-like thickened just beyond hypandrium; slightly sinuate, with broad even curvature shortly beyond
basal thickening, gently curved on remaining part. Ejaculatory apodeme not extended beyond basal curvature of
phallus, with broad lateral wings and similar vertical wing.
Female. Unknown.
Distribution. Eurasia; known only from Wrangel Island.
Etymology. The specific name refers to the similarity of the new species to R. macrura Loew.
Remarks. The new species belongs to the R. pusilla group sensu Barták & Kubík (2009) (see also Sinclair et
al. 2019: 65). It resembles R. macrura Loew described from East Siberia (Irkutsk) (Loew 1871). Rhamphomyia
macrura is somewhat smaller (wing length 4.8 mm vs. 6.7 mm in R. submacrura sp. nov.), has brush-like arranged
long setae on the mid basitarsus and much longer epandrial lamellae (Frey 1954: 427, textfig. 214), which are nearly
as long as the abdomen. The females of both species are unknown.
Rhamphomyia (Pararhamphomyia) subtenuiterfilata Shamshev & Sinclair sp. nov.
(Figs 45–47)
urn:lsid:zoobank.org:act:A3936415-AFA7-4469-8A77-E07648EAAAF3
Type material. HOLOTYPE male, labelled: [Chukotka AO (Wrangel Island):] “[printed in Cyrillic, Russian]
bass. [=basseyn, basin] r. [=reka, river] Khischniki/ 7 km SO g. [=gora, mountain] Sovetskaya/ o. [=ostrov, island]
Wrangel/ Gorodkov 15.vii.[1]972”; “Holotypus/ Rhamphomyia/ subtenuiterfilata/ Shamshev, Sinclair sp. n. [red
label]” (INS_DIP_0000616, ZIN; terminalia dissected, in microvial pinned with specimen).
PARATYPES: RUSSIA. Chukotka AO (Wrangel Island): Somnitelnaya Bay, 400 m, S Sovetskaya Mtn,
20.vii.1966, KBG (1 ♀, ZIN). Yakutia: Indigirka River, lower flow of river Ystan-Yuryakh, Momskiy District,
29.vi.1976, V. Kovalev (1 ♂, ZMMU).
Diagnosis. Mid-sized (wing length about 4.5 mm), blackish, subshiny flies. Male holoptic; thorax and abdomen
black setose, scutum subshiny, anepisternum pruinescent; legs dark brown, hind basitarsus clavate, hind tibia with
moderately long setae and hind tarsomeres 1–2 with numerous long fine setae dorsally; abdomen almost shiny,
epandrium and cerci elongate, cercus spatulate beyond dorsal cavity; phallus well exposed, filamentous, long, gently
curved. Female with brownish infuscate wing, short setose slender legs without pennate setae.
Description. Wing length 4.7 mm. Male (Fig. 45). Head with faint greyish pruinescence on face, frons,
postgena, ocellar triangle and occiput; dark setose. Holoptic; upper ommatidia enlarged. Frons represented by very
small subtriangular space below ocellar tubercle and larger subtriangular space above antennae, bare. Face broad,
bare. Ocellar triangle with 2 moderately long fine and some short setae. Occiput with numerous long fine setae,
including postoculars; postgena with numerous hair-like setae. Antenna blackish; scape short, slightly longer than
pedicel, both with short setulae; postpedicel conical, nearly 2X longer than wide; stylus very short, slightly shorter
than half of postpedicel basal width. Palpus dark; with several long, dark fine setae. Proboscis with labrum dark
brown, nearly as long as head height.
Thorax dark in ground-colour, faintly greyish pruinescent (except noted), with black setation; postpronotal lobe
and mesonotum almost uniformly shiny, very faintly pruinescent (somewhat denser on scutellum). Proepisternum
with few moderately long fine setae on lower section; upper proepisternum in front of spiracle without setae.
Prosternum bare. Antepronotum with several short setae. Postpronotal lobe with 1 long stronger and several shorter
fine setae. Mesonotal setation: 1 presut spal (with some additional short fine setae), 3 npl (with several additional
fine setae anteriorly), 1 psut spal (with additional setulae anteriorly), 1 pal, 4 sctl (in holotype 5); acr very short,
cilia-like, arranged in 2 close irregular rows, lacking on prescutellar depression; presutural dc similar to acr, fine,
arranged in 2 irregular rows, postsutural dc 1–2-serial, 3–4 prescutellars longest. Laterotergite with several long fine
setae. Anterior and posterior spiracles brown.
SHAMSHEV ET AL.
44 · Zootaxa 4848 (1) © 2020 Magnolia Press
FIGURES 45–47. Rhamphomyia (Pararhamphomyia) subtenuiterfilata sp. nov., male, holotype. 45. habitus, lateral view; 46.
terminalia, lateral view; 47. cerci, dorsal view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium;
hypd—hypandrium; ph—phallus.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 45
Legs slender (except noted), uniformly brownish; mostly shiny, coxae and trochanters faintly greyish
pruinescence; black setose. Fore coxa covered with long fine setae anteriorly. Hind femur slightly thickened on
about apical half; hind tibiae slightly evenly thickened toward apex; hind basitarsus clavate, about 1.5X broader at
middle than apex of hind tibia; hind tarsomere 2 expanded. All femora whitish pilose ventrally, short setose (except
noted). Fore femur with minute anteroventral and posteroventral setae. Mid femur with rows of moderately long
anteroventrals (4–5 setae near base longer) and posteroventrals (somewhat longer on about apical half). Hind femur
with numerous moderately long setae anteroventrally on about apical third. Fore and mid tibiae with some slightly
longer fine setulae posteriorly, mid tibia with numerous stronger long anteroventral and posteroventral setae on
about apical 2/3; hind tibia with moderately long fine setae dorsally (longest setae only slightly longer than tibia
width), 1 long seta in posteroapical comb. Fore and mid tarsomeres short setose; hind tarsomeres 1–2 with long,
dense erect fine setae dorsally, tarsomere 3 with slightly longer setae dorsally.
Wing membrane faintly infuscate, with brownish veins; CuA+CuP (anal vein) incomplete, evanescent apically,
almost reaching wing margin as fold. Pterostigma brownish yellow. Basal costal seta absent. Anal lobe well-
developed; axillary incision right-angled. Squama brown, dark fringed. Halter brownish.
Abdomen dark brown, almost shiny, very faintly pruinescent; mostly with black fine moderately long setae.
Segments 6–7 unmodified. Segment 8 with separated tergite and sternite; tergite 8 simple, entire, subtriangular
viewed laterally, very narrow mid-dorsally, broadly concave anteriorly, with almost straight posterior margin,
numerous moderately long setae along posterior margin; sternite 8 simple, enlarged, subrectangular in lateral view,
V-shaped viewed posteriorly, covered with numerous long setae posteriorly.
Terminalia (Fig. 46) dark brown, directed obliquely upwards. Cerci (Fig. 47) separated from each other and
from epandrium, not extended beyond tergite 8 anteriorly; cercus long, broadly deeply concave on upper margin
about middle, its middle portion very narrow, apical portion spatulate (lateral view), about 2.5X as broad as middle
portion, with small internal subrectangular projection slightly beyond middle of cercus (dorsal view); covered with
short fine setae on broadened anterior and posterior parts, lacking setation on narrowed middle portion; without pair
of subepandrial lobes beneath. Epandrium subrectangular, elongate, about 4X as long as broad, broadly concave on
upper margin, with straight lower margin, somewhat broadened and angularly expanded apically, extended slightly
beyond cerci posteriorly; with row of 6–7 long dark setae near upper anterior corner, covered with dense fine setulae
on broadened apical part, some numerous moderately long setae along lower margin. Hypandrium very narrow,
rim-like, entire, well sclerotized; bare. Phallus well exposed, long; mostly hair-like, knob-like thickened just beyond
hypandrium; with right-angle curvature shortly beyond basal thickening, otherwise gently sinuate. Ejaculatory
apodeme slightly extended beyond basal curvature of phallus, with narrow lateral wings and broad vertical wing.
Female. Frons broad, greyish pruinescent, with marginal setulae; body with shorter setation; hind femur
uniformly slender, with short anteroventral setae over entire length; hind tibia and tarsi slender, covered with short
setae; wing darker, distinctly brownish infuscate; abdominal segments 5–8 somewhat denser pruinescent (lateral
view); cercus long, slender, with dark setulae.
Distribution. The new species is currently known only from Yakutia and Wrangel Island of Russia.
Etymology. The specific name refers to the similarity of the new species to R. tenuiterfilata Becker, 1900.
Remarks. The new species is very similar to R. tenuiterfilata (known from Taymyr Peninsula) in the structure
of the male hind leg and the terminalia. However, R. subtenuiterfilata differs from R. tenuiterfilata primarily by
brownish halteres (vs. yellowish) and black setose abdomen (vs. pale setose).
Habitat. On Wrangel Island, the species is known from single specimens collected in the mountains.
Rhamphomyia (Pararhamphomyia) ursinella Melander
(Fig. 48)
Rhamphomyia ursina Malloch, 1919: 46. Type-locality: Bernard Harbour, Nunavut, Canada.
Rhamphomyia ursinella Melander, 1928: 209 (replacement name for ursina Malloch, preoccupied by Oldenberg, 1915: 91).
Other references: Sinclair et al., 2019: 71 (type material, redescription, full list of references).
Material examined. RUSSIA. Yakutia (New Siberian Islands): centre of Kotelny Island, upper flow of river
Balykhtakh near mouth of river Tuguttakh, arctic desert on flowers, 15.vii.1973, KBG (20 ♂, 20 ♀, ZIN).
Recognition. Mid-sized (wing length 3.6–4.5 mm) blackish flies. Male (Fig. 48) holoptic; anepisternum with
SHAMSHEV ET AL.
46 · Zootaxa 4848 (1) © 2020 Magnolia Press
shiny spot; scutum with 2 narrow shiny median vittae (sometimes with less distinct subshiny vitta on supra-alar
space); thorax black setose, presutural dc long, fine, 2–3-serial, acr similar to dc, biserial, 10–12 sctl; legs dark
brown, hind basitarsus thickened apically, all tibiae and basitarsi clothed in dense long fine setae, hind tibia covered
with these setae over entire dorsal face; wing infuscate, CuA+CuP incomplete, halter brown; abdomen very faintly
pruinescent, almost shiny, with black setae; epandrium elongate, slightly constricted at middle; cercus elongate,
about 2/3 length of epandrium, slightly constricted medially; phallus well exposed, filamentous, long, gently curved
slightly beyond epandrium. Female with darker wing, short setose slender legs without pennate setae.
Distribution. Holarctic; R. ursinella is widespread across the low to high arctic region of North America, east
of Hudson Bay (Sinclair et al. 2019). In Eurasia, this species was found only from New Siberian Islands of Arctic
Russia.
FIGURE 48. Rhamphomyia (Pararhamphomyia) ursinella Melander, male habitus, lateral view.
Rhamphomyia (Pararhamphomyia) wrangeli Shamshev & Sinclair sp. nov.
(Figs 49–53)
urn:lsid:zoobank.org:act:D1A835B2-FA2B-4104-B593-F6A43BDDFA89
Type material. HOLOTYPE ♂, labelled [RUSSIA. Chukotka AO (Wrangel Island)]: “[printed in Cyrillic,
Russian] bukh. [=bukhta, bay] Somnitelnaya,/ south of o. [=ostrov, island] Wrangel/ Gorodkov 19.vii.[1]966”; “on
flower/ Dryas integrifolia,/ spotted/ tundra”; “Holotypus/ Rhamphomyia/ wrangeli/ Shamshev, Sinclair sp. n. [red
label]” (INS_DIP_0000617, ZIN).
PARATYPES: RUSSIA: Chukotka AO (Wrangel Island): same data as holotype, (1 ♂, 7 ♀, ZIN; 1 ♂, 1
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 47
♀, CULSP); environs of Somnitelnaya Bay, south of Wrangel Island, valley of river Somnitelnaya, Mineev Mtns,
150 m, pebbles, on flower of Dryas integrifolia, 19.vii.1966, KBG (1 ♂, 4 ♀, ZIN; 1 ♂, 1 ♀, CNC); same locality,
on flower of Potentilla emarginata, 22.vii.1966, KBG (2 ♂, ZIN).
Diagnosis. Small dark flies (wing length about 3.5 mm). Male holoptic; scutum almost uniformly brownish
grey pruinescent, acr and dc biserial, usually 6 sctl; wing faintly brownish, anal vein incomplete, halter brown;
hind tibia strongly curved inward on about basal third; abdominal segments 7 and 8 asymmetrical; terminalia small,
slightly rotated to right, phallus mostly hidden. Female legs with simple setae.
Description. Body length 3.2–3.5 (holotype 3.3); wing length 3.2–3.4 (holotype 3.2) mm. Male (Fig. 49). Head
with dense greyish brown pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Holoptic;
upper ommatidia enlarged. Frons represented by very small subtriangular space below ocellar tubercle and larger
subtriangular space above antennae, bare. Face broad, bare. Ocellar triangle with several long fine setae. Occiput
with numerous long setae, including postoculars; postgena with numerous hair-like setae. Antenna blackish brown;
scape short, slightly longer than pedicel, both with short setulae; postpedicel conical, with slightly concave ventral
margin on apical part, nearly 2.5X longer than wide; stylus short, nearly as long as postpedicel basal width. Palpus
dark; with several long, dark fine setae. Proboscis with labrum dark reddish-brown, nearly as long as head height.
Thorax dark in ground-colour, greyish brown pruinescent; with black setation; scutum almost uniformly
brownish grey pruinescent (dorsal view), without vittae. Proepisternum with tuft of several moderately long fine
setae on lower section; upper proepisternum in front of spiracle without setae. Prosternum bare. Antepronotum with
several short setae. Postpronotal lobe with 1 long and several short setae. Mesonotum with well-differentiated setae
but their number, position and robustness variable; 1 presutural intra-alar, 1–2 presut spal (with some additional
short fine setae), 3–4 npl (with finer short setae anteriorly), 1–2 psut spal (with additional setulae anteriorly), 2 pal,
6–8 sctl (usually 6); acr short, arranged 2 close irregular rows, lacking on prescutellar depression; presutural dc
longer than acr, arranged in 2 irregular rows, postsutural dc 1–2-serial, 3–4 prescutellars longest. Laterotergite with
numerous long fine setae. Anterior and posterior spiracles brown.
Legs slender, uniformly brownish; mostly subshiny, coxae and trochanters with denser greyish pruinescence;
black setose. Fore coxa covered with long fine setae anteriorly. Hind tibia strongly curved inward on about basal
third (posterior view), remaining podomeres unmodified. Mid and hind femora whitish pilose ventrally. Fore femur
with rows of anteroventral and posteroventral very short fine setae (somewhat longer near base of femur), some
moderately long setae anteriorly toward apex; mid femur with rows of short anteroventral and posteroventral setae
(longest setae at most as long as half of femur width), some moderately long setae anteriorly; hind femur with
moderately long anteroventral setae somewhat denser and longer on about apical third, some similar setae anteriorly
and posteriorly. Fore tibia covered with moderately long fine setae posterodorsally (some setae usually slightly
stronger); mid tibia with 3–4 posterodorsal, short spine-like setae ventrally toward apex; hind tibia covered with
moderately long setae dorsally (longest setae slightly longer than tibia median width); no seta in posteroapical
comb. Fore basitarsus with similar to fore tibia setae posterodorsally; hind basitarsus with similar to hind tibia setae
posterodorsally.
Wing membrane faintly brownish, with brownish veins; CuA+CuP (anal vein) incomplete. Pterostigma brownish
yellow. One basal costa seta present. Anal lobe well-developed; axillary incision right-angled. Squama brown, dark
fringed. Halter brown.
Abdomen (Fig. 50) with segments 1–6 brownish, segments 7–8 blackish brown, all segments faintly brownish
grey pruinescent, subshiny; segments 1–6 mostly covered with moderately long dark setae, short setae on tergites
dorsally. Tergite and sternite 6 slightly concave posteriorly. Sclerites of segments 7 and 8 strongly modified (Figs
51–53), asymmetrical and stronger sclerotized (darker) than preceded segments. Segment 7 with tergite and sternite
narrowly fused anteriorly. Tergite 7 slightly asymmetrical, large, longer than tergite 6, subrectangular viewed
dorsally, with straight posterior margin; minute setulae posteriorly. Sternite 7 strongly asymmetrical, inflated,
narrowly deeply concave posteromedially, with produced left lateral margin, free laying internal lateral narrow
sclerite supporting anterior lateral projections of segment 8; covered with numerous short setae denser and somewhat
stronger around posterior concavity. Segment 8 slightly rotated to right, with fused tergite and sternite (no trace of
suture), slightly asymmetrical ventrally; large subtriangular anterior dorsomedial projection and 2 broad digitiform
anterior lateral projections inserted into tergite 7, subtriangular anterior ventromedial projection (corresponding
to posterior concavity on sternite 7); numerous long setae along posterior margin laterally and ventrally, scattered
minute setulae dorsally.
SHAMSHEV ET AL.
48 · Zootaxa 4848 (1) © 2020 Magnolia Press
FIGURES 49–53. Rhamphomyia (Pararhamphomyia) wrangeli sp. nov., male. 49. habitus, holotype, lateral view; 50. abdomen,
right lateral view; 51. postabdomen, macerated, right lateral view; 52. same, macerated, dorsal view; 53. same, outline (setation
omitted), dorsal view. Abbreviations: cerc—cercus; epand—epandrium; ph—phallus; sg 8—segment 8; st 7—sternite 7; tg
7—tergite 7.
Terminalia (Figs 51–53) slightly rotated to right, small, dark brown. Cerci separated from epandrium, not
extended anteriorly beyond posterior margin of tergite 8, without pair of subepandrial lobes beneath; in lateral view
cercus shorter than epandrium, subrectangular, with straight upper margin, slightly rounded apically; in dorsal view
cercus strongly convex laterally and deeply concave on inner margin, with short broad subapical projection directed
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 49
downward; mostly covered with minute setulae, subapical projection with dense spinule-like setulae. Epandrium
narrowly subtriangular; with some moderately long setae along lower margin and closer to apex. Hypandrium
invisible in situ, mostly membranous, only lateral arms sclerotised. Phallus mostly hidden, only short portion
between posterior margin of sternite 8 and lower margin of epandrium visible; short, thicker on about basal third
and uniformly slender on remaining portion, gently arcuate, hook-like curved to left at extreme apex. Ejaculatory
apodeme slightly extended beyond basal curvature of phallus, with subequally narrow lateral and vertical wings.
Female. Similar to male, except frons broad, with marginal setulae, ommatidia of equal size; sometimes upper
proepisternum in front of spiracle with 1 seta; legs with short setae, no pennate setae, hind tibia simple; wing darker,
uniformly brownish infuscate; cercus long slender, with dark setulae.
Distribution. Russia (Chukotka Autonomous Okrug: Wrangel Island).
Etymology. The specific name is a patronym in honour of Ferdinand Petrovich Wrangel (Baron Ferdinand von
Wrangel, a Baltic German), a Russian explorer and seaman, and one of the founders of the Russian Geographic
Society. Wrangel Island was named in 1867 by Thomas W. Long, an American whaling captain.
Remarks. The male of the new species can be readily distinguished from other groups of Pararhamphomyia
recognised by Barták & Kubík (2009) in the characters noted in the key. Several undescribed species of this group
occur in North America.
Habitat. This species was collected from Wrangel Island only in 1966, and in one area (environs of Somnitelnaya
Bay), mainly on the flowers of Dryas integrifolia.
Rhamphomyia (Rhamphomyia) albosegmentata (Zetterstedt)
(Fig. 54)
Rhamphomyza albosegmentata Zetterstedt, 1838: 563. Type locality: “Lapponia Umensi, Stoettingsfjellet inter Lycksele et
Åsele”, Sweden.
Other references: Barták, 1981: 367 (holotype data).
Rhamphomyza arctica Zetterstedt, 1838: 566. Type locality (by lectotype designation): “in inferalpinis Finmarkiae occident, ad
Bossekop; E Lappon. Tornensi”, Norway.
Other references: Zetterstedt, 1842: 393 (synonymy); Collin, 1961: 401 (type data); Barták & Danielson, 2007: 107 (lectotype
designation).
Rhamphomyza engadinica Oldenberg, 1922: 344. Type locality (by lectotype designation): Oberengadin, Swiss Alps.
Other references: Barták, 1981: 368 (lectotype designation, synonymy).
Rhamphomyia (Alpinomyia) albosegmentata (Zetterstedt): Frey, 1956: 557 (revision).
Rhamphomyia (Rhamphomyia) albosegmentata (Zetterstedt): Frey, 1922: 74 (key); Collin, 1961: 399 (revision); Gorodkov &
Kovalev, 1969: 635 (key); Barták, 1981: 367 (revision); 1982: 396 (revision); Chvála & Wagner, 1989: 282 (catalogue);
Yang et al., 2007: 179 (catalogue); Shamshev, 2016: 72 (checklist).
Material examined. RUSSIA. Murmanskaya Prov. (Kildin Island): Kola Bay, 23.viii.–1.ix.1923, Serebrjansky
(1 ♂, ZIN). Nenets AO (Kolguev Island): Bugrino, sea shore, meadow, 21–22.viii.1970, KBG (3 ♂, 2 ♀, ZIN).
Recognition. Mid-sized (wing length 5–6.5 mm) greyish, dark-legged, black setose flies. Male (Fig. 54):
holoptic; antennal postpedicel and stylus black, scape and pedicel somewhat paler; labrum 1.1–1.5X longer than
head height; palpus black, long, with several long strong setae. Thorax grey pruinose, scutum somewhat finer
pruinescent, without vittae; prosternum bare, upper proepisternum in front of spiracle with a few fine setae;
mesonotal setae strong bristle-like (including acr and dc), acr biserial, presutural dc 2–3-serial, 4–6 sctl; legs
blackish brown, long, slender, mostly shiny to subshiny (except coxae); fore femur with single row of very long
posteroventral setae; hind femur with 3–4 anterodorsal setae on about apical half, 3–6 strong anteroventral setae of
different lengths over entire femur length and 1 long strong posteroventral seta on about basal third; all tibiae with
rows of numerous very long strong setae dorsally, hind tibia with 1 long seta in posteroapical comb; fore basitarsus
with several long setae dorsally. Wing hyaline, CuA+CuP (anal vein) complete, entirely sclerotized, 1 long basal
costal seta present, axillary incision very acute, halter brown. Abdomen (except terminalia) densely uniformly light
grey pruinescent, tergites with very long posteromarginal setae laterally; segment 7 with unmodified structure and
vestiture; segment 8 with tergite and sternite separated; tergite 8 upturned posteriorly, with slightly convex posterior
margin, without projections; sternite 8 nearly as long as sternite 7. Terminalia with cerci and epandrium brownish
to reddish brown, faintly pruinescent; cerci separated from each other and from epandrium, not extended anteriorly
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50 · Zootaxa 4848 (1) © 2020 Magnolia Press
beyond tergite 8; cercus slightly produced anteriorly, prolonged slightly beyond epandrium middle posteriorly, with
narrow cavity near base dorsally forming short finger-like basal projection, rather trapezoid-like beyond cavity,
lower posterior corner slightly produced downward, mostly covered with minute setulae, lower posterior corner
with dense spinules, without pair of subepandrial lobes beneath; epandrium subtriangular, narrow, rounded apically,
with long to moderately long setae more numerous along lower margin; subepandrial sclerite divided; hypandrium
yellow, shiny, subtriangular, bare; phallus thick, vesicular apically; ejaculatory apodeme extended far beyond
basal curvature of phallus, without lateral wings. Female: similar to male except, eyes dichoptic, frons very broad,
parallel-sided; femora mostly with short setulae, only hind femur with 2–4 strong anteroventral setae; wing faintly
brownish infuscate; abdominal tergites 5–8 and sternites with shorter setation.
Distribution. Palaearctic; Fennoscandia, Great Britain, mountains of Central Europe, across northern Russia
from Murmanskaya Province to Taymyr Peninsula.
Rhamphomyia (Rhamphomyia) morio (Zetterstedt)
(Fig. 55)
Rhamphomyza morio Zetterstedt, 1838: 565. Type locality (by lectotype designation): Alteidet, Norway.
Other references: Collin, 1961: 399 (type data); Barták, 1981: 395 (lectotype designation).
Rhamphomyza villosa Zetterstedt, 1838: 563. Type locality: “Dowrefjell Norvegiae (Lappon. Norvegica)”, Norway.
Other references: Collin, 1961: 399 (type data); Barták, 1981: 396 (holotype data).
Rhamphomyza fumipennis Zetterstedt, 1859: 5019. Type locality: “Lapponia Umensi, ad Tarna”, Sweden.
Other references: Collin, 1961: 399 (type data); Barták, 1981: 396 (holotype data).
Rhamphomyia (Alpinomyia) morio (Zetterstedt): Frey, 1956: 573 (revision).
Rhamphomyia (Rhamphomyia) morio (Zetterstedt): Frey, 1922: 77 (key); Collin, 1961: 397 (revision); Gorodkov & Kovalev,
1969: 635 (key); Barták, 1981: 395 (revision), 1982: 408 (revision); Chvála & Wagner, 1989: 287 (catalogue); Yang et al.,
2007: 185 (catalogue); Shamshev, 2016: 78 (checklist).
Material examined. RUSSIA. Murmanskaya Prov. (Kildin Island): 13.vii.1908, L. Bianki (1 ♂, ZIN).
FIGURES 54, 55. Rhamphomyia (Rhamphomyia), male, lateral view. 54. R. (R.) albosegmentata (Zetterstedt); 55. R. (R.)
morio (Zetterstedt).
Recognition. Large (wing length 6–7.5 mm) blackish grey, dark-legged, black setose flies. Male (Fig. 55):
holoptic, eyes not quite touching; antenna black, scape nearly 3X longer than pedicel, both with long setae; labrum
about 1.5X longer than head height; palpus black, long, with numerous long fine setae; thorax mostly densely greyish
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 51
pruinescent, scutum greyish brown, somewhat lustrous; prosternum bare, upper proepisternum in front of spiracle
with several long fine setae; mesonotal setae intermixed with additional long fine setae, often hardly differentiated,
6–8 sctl; acr long, fine, 2–3-serial, presutural dc slightly longer than acr, 3–4-serial, not separated by bare space
from supra-alars anteriorly; legs black, long, slender, mostly shiny to subshiny (except coxae); fore femur covered
with dense, very long, fine, irregularly spread setae posteroventrally; hind femur with several anterodorsal setae on
about apical half, 4–8 strong short anteroventral setae over entire femur length and some similar setae posteriorly,
only pale pilose ventrally; all tibiae with rows of numerous very long strong setae dorsally, hind tibia with 1 long
seta in posteroapical comb. fore and hind basitarsi with some longer setae dorsally; wing faintly brownish infuscate,
CuA+CuP (anal vein) complete, entirely sclerotized, 1 long basal costal seta present, axillary incision very acute,
halter brown; abdomen uniformly light grey pruinescent, denser on sternites; tergites somewhat lustrous, with very
long posteromarginal setae; terminalia (Barták 1981: 396, fig. 17a) very similar to R. albosegmentata.
Distribution. Palaearctic; Fennoscandia, Great Britain, mountains of Central Europe, northern European
Russia.
Species unplaced to subgenus
Rhamphomyia hirtula Zetterstedt
(Fig. 56)
Rhamphomyia hirtula Zetterstedt, 1842: 421. Type-locality: Greenland.
Other references: Barták & Danielson, 2007: 111 (type data); Sinclair et al., 2019: 78 (revision, redescription, full list of
references).
Rhamphomyia (Pararhamphomyia) hirtula Zetterstedt: Frey, 1955b: 482 (revision); Chvála & Wagner, 1989: 301 (catalogue);
Yang et al., 2007: 171 (catalogue); Shamshev, 2016: 64 (checklist).
Rhamphomyia (Rhamphomyia) hirtula Zetterstedt: Collin, 1961: 401 (revision).
Material examined. RUSSIA. Archangelskaya Prov. (Novaya Zemlya Archipelago): Novaja Zemlja, acad.
Baer, 1837 (2 ♂, ZIN); Propaschaya Guba, sea shore, in mountains, 18.vii.1912, S. Skribova (1 ♀, ZIN).
Recognition. Mid-sized (wing length 5.5–6 mm) blackish, dark-legged, black setose robust flies. Male (Fig.
56): holoptic; antenna dark, scape and pedicel with long fine setae on outer side, minute setulae on inner side; palpus
dark with numerous long fine setae; labrum nearly 1.3X longer than head height; thorax mostly greyish brown
pruinose; scutum dark brown pruinescent, slightly lustrous, with two hardly visible and slightly paler vittae between
rows of acr and dc; prosternum bare, upper proepisternum in front of spiracle with several long fine setae; mesonotal
setae mostly fine (including acr and dc), hardly differentiated, acr 2–3-serial, present on anterior half of prescutellar
depression, presutural dc multiserial, not separated from supra-alars by bare space, 12–20 sctl; legs robust, hind
tibia slightly uniformly incrassate; fore femur with numerous long fine setae anteroventrally, posteroventrally and
posteriorly; mid femur with very short anteroventral (except near base) and somewhat longer posteroventral setae;
hind femur with numerous moderately long setae anteroventrally and ventrally; fore and mid tibiae with rows of
several moderately long strong setae dorsally and some fine setae posteriorly; hind tibia with numerous long strong
setae dorsally and long fine setae anteroventrally, 1 short seta in posteroapical comb; fore basitarsus with some
short fine setae posteriorly, hind tarsomeres 1–3 with several long setae dorsally; wing faintly infuscate, CuA+CuP
(anal vein) complete, entirely sclerotized, 1 long basal costal seta present, axillary incision acute, halter with brown
knob and reddish brown stem; abdomen densely uniformly greyish brown pruinescent, with numerous long setae;
structure of segments 7–8 as in R. kaninensis but posteromarginal setae more numerous and stronger; terminalia
almost identical to R. kaninensis (including shape of phallus) but larger, cercus more gently rounded apically, with
broader basal projection, epandrium with more numerous strong setae and denser cluster of fine setae subapically.
Female similar to male, except eyes dichoptic, frons very broad, parallel-sided, body (especially legs and abdomen)
with much shorter setation; wing slightly darker, faintly brownish infuscate; legs without pennate setae, hind tibia
somewhat dilated, almost bare anteriorly.
Distribution. Holarctic; R. hirtula appears to have a disjunct distribution. In North America, this species is
recorded from several localities in Yukon and Greenland; in Eurasia, it is known from Iceland and Scotland, as
well as from Novaya Zemlya Archipelago, Taymyr, Chukotka and high mountains of Altay Republic of Russia
(Shamshev & Barkalov 2009; Sinclair et al. 2019).
SHAMSHEV ET AL.
52 · Zootaxa 4848 (1) © 2020 Magnolia Press
Remarks. Shamshev (2016: 64) has noted R. hirtula from Wrangel Island, but with a question mark. Our study
did not confirm the presence of this species on this island. Rhamphomyia hirtula belongs to the R. hirtula group.
Rhamphomyia kaninensis Frey
(Figs 57–61)
Rhamphomyia kaninensis Frey in Lundström & Frey, 1913: 7. Type locality: Arkhangelskaya Province, “Kambalnitsa” [~67°52′N
44°09′E], Russia.
Other references: Frey, 1922: 42 (key), 1955b: 483 (revision); Gorodkov & Kovalev, 1969: 624 (key); Chvála & Wagner, 1989:
286 (catalogue); Shamshev, 2001b: 333 (key); Yang et al., 2007: 184 (catalogue); Shamshev, 2016: 77 (checklist).
Material examined. RUSSIA. Krasnoyarskiy Terr. (Dikson Island): small island “Konus”, 31.vii.1948,
Korotkevich (1 ♂, ZIN).
Additional material. RUSSIA. Krasnoyarskiy Terr. (Taymyr Peninsula): Engelgardt Lake, NE shore,
valley of brook, 2.vii.1967, KBG (3 ♂, 5 ♀, ZIN; 1 ♂, 1 ♀, CNC).
Diagnosis. Mid-sized (wing length about 4.5 mm), robust, black setose flies with brownish thorax and greyish
abdomen. Male mid tibia and mid basitarsus with numerous very long setae dorsally; hind tibia slightly uniformly
thickened, hind basitarsus slender; abdomen densely light grey pruinescent; wing whitish, CuA+CuP (anal vein)
complete, halter brown; abdominal tergite 8 without projections; cercus with finger-like projection near base
dorsally; phallus well exposed, slender, gently sinuate on about middle. Female wing uniformly faintly brownish
infuscate, legs without pennate setae.
Redescription. Body length 4.5–4.9; wing length 4.3–4.5 mm. Male (Fig. 57). Head with dense greyish brown
pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Holoptic; upper ommatidia enlarged;
eyes touching. Frons represented by very small subtriangular space below ocellar tubercle and larger subtriangular
space above antennae, bare. Face broad, bare. Ocellar triangle with 2 long and several shorter fine setae. Occiput
with numerous long fine setae, including postoculars; postgena with numerous hair-like setae. Antenna blackish
brown; scape slightly longer than pedicel, both with somewhat longer fine setae on outer side and short setulae on
inner side; postpedicel conical, nearly 3X longer than wide; stylus short, nearly as long as postpedicel basal width.
Palpus dark, long; with numerous long, dark fine setae. Proboscis with labrum dark reddish-brown, nearly 1.3X as
long as head height.
Thorax dark in ground-colour, mostly greyish brown pruinescent; with black setation; scutum almost uniformly
brownish pruinescent (dorsal view), slightly lustrous, only some traces of two narrow dark vittae between rows
of acr and dc. Proepisternum with tuft of several long fine setae on lower section; upper proepisternum in front of
spiracle with 3–5 similar setae. Prosternum bare. Antepronotum with numerous short setae. Postpronotal lobe with
1 (sometimes 2) longer somewhat stronger and numerous shorter fine setae. Mesonotal setae generally quite well
differentiated (number, position and robustness variable); 1–2 presut spal (with additional shorter setae), 3–4 npl
(with additional shorter setae anteriorly), 1–2 psut spal (with additional setae anteriorly), 1 long and 1 short pal, 4–6
sctl; acr moderately long, fine, arranged in 2 irregular rows, lacking on prescutellar depression; presutural dc slightly
longer than acr, 3–4-serial, hardly separated by from supra-alar setae, postsutural dc 1–2-serial, 3–4 prescutellars
longest. Laterotergite with numerous long fine setae. Anterior and posterior spiracles brown.
Legs robust, uniformly dark brown; mostly subshiny, coxae and trochanters with denser greyish pruinescence;
black setose. Fore coxa covered with long fine setae anteriorly. Hind tibia slightly uniformly thickened. Femora whitish
pilose ventrally. Fore femur covered with very short setae including rows of fine anteroventrals and posteroventrals.
Mid femur with similar setation to fore femur but anteroventral and posteroventral setae somewhat longer. Hind
femur mostly with very short setae anteroventrally, only 6–7 anteroventral setae on about apical third moderately
long; covered with moderately long setae anteriorly and on basal part posteriorly. Fore tibia with numerous fine,
moderately long setae over entire length posterodorsally, 2–3 similar anterodorsal setae on about basal third. Mid
tibia (Fig. 58) with numerous dense very long (2–2.5X longer than tibia width) setae dorsally and posterodorsally;
numerous anteroventral and posteroventral setae becoming spine-like closer to apex. Hind tibia (Fig. 59) with
numerous long setae dorsally and shorter setae anteroventrally; 1 seta in posteroapical comb. Fore basitarsus with
some fine setae posterodorsally (as on fore tibia); mid basitarsus with several very long setae dorsally (as on mid
tibia); hind basitarsus with several long setae dorsally.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 53
Wing membrane whitish, with brownish veins; all veins complete (except Sc), CuA+CuP (anal vein) weaker
sclerotised on about basal half. Pterostigma brownish yellow. Basal costal seta present, long. Anal lobe well-
developed; axillary incision right-angle. Squama brownish, dark fringed. Halter with brown knob and reddish
brown stem.
FIGURES 56–61. Rhamphomyia, male. 56. R. hirtula Zetterstedt, habitus, lateral view; 57–61. R. kaninensis Frey. 57. habitus,
lateral view; 58. mid leg, anterior view; 59. hind leg, anterior view; 60. terminalia, lateral view; 61. phallus, lateral view.
Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium; hypd—hypandrium; ph—phallus.
Abdomen dark, densely light grey pruinescent; covered with short to moderately long black fine setae (except
noted). Tergite 7 with unmodified structure but bearing 4 erect medial posteromarginal setae longer and stronger than
posteromarginal setae on tergite 6. Sternite 7 with similar structure and setation to sternite 6. Segment 8 with tergite
SHAMSHEV ET AL.
54 · Zootaxa 4848 (1) © 2020 Magnolia Press
and sternite separated but pleural space indistinct. Tergite 8 about 2X shorter than tergite 7, somewhat upturned
posteriorly, with truncate posterior margin, without projections; bearing 2 long, strong erect posteromarginal setae
medially. Sternite 8 large, somewhat longer than sternite 7 (lateral view); with numerous dense very long setae near
posterior margin.
Terminalia (Figs 60, 61) brownish, black setose. Cerci separated from each other and from epandrium, not
extended anteriorly beyond tergite 8; covered with mostly fine setulae, some marginal setulae on inner face stronger;
cercus slightly shorter than epandrium, subrectangular, rounded apically (lateral view), with dorsal notch forming
long pointed finger-like projection on about basal third; short broad inner projection subapically (dorsal view);
without pair of subepandrial lobes beneath. Epandrium subtriangular, rounded apically, with several long setae
on about middle and along lower margin, bearing cluster of somewhat finer numerous dense setae near apex.
Hypandrium invisible in situ, rim-like, entirely sclerotized, bare. Phallus (Fig. 59) exposed on about middle third;
expanded on basal part, gently sinuate near middle, not extended beyond cerci. Ejaculatory apodeme slightly
extended beyond basal curvature of phallus, with broad lateral wings and somewhat narrower vertical wing.
Female (described for first time). Similar to male, except frons very broad, parallel-sided, with marginal setulae;
eye ommatidia equally small; ocellar tubercle and occiput with shorter and stronger setae; thorax with somewhat
shorter setation; legs covered with short simple setae; wing membrane uniformly faintly brownish infuscate, anal
vein uniformly sclerotised; abdomen slightly darker pruinescent; cercus long slender, with dark setulae.
Distribution. Holarctic; in Eurasia the species is known only from the arctic region of Russia. This species is
also known from North America (T. Saigusa, unpubl. data), but further details were not available.
Remarks. Rhamphomyia kaninensis belongs to the R. hirtula group. Frey described this species after three
males collected from the Kanin Peninsula (Arkhangelskaya Province, Russia), one male in MZH was examined by
IVS. Frey (1913: 8) noted that this species was collected on bushes of Salix in the tundra.
Rhamphomyia sublongiseta Shamshev, Sinclair & Saigusa sp. nov.
(Figs 62–65)
urn:lsid:zoobank.org:act:99F127D3-A192-4D59-9030-EEEB1F7E3827
Type material. HOLOTYPE ♂, labelled: [RUSSIA, Chukotka AO (Wrangel Island)] “[printed in Cyrillic,
Russian] bukh. [=bukhta, bay] Somnitelnaya/ south of o. [=ostrov, island] Wrangel/ Gorodkov 19.vii.[1]966”; “on
flower/ Dryas integrifolia,/ spotted/ tundra”; “Holotypus/ Rhamphomyia/ sublongiseta Shamshev,/ Sinclair, Saigusa
sp. n. [red label]” (INS_DIP_0000618, ZIN).
PARATYPES: CANADA. Yukon: km 155 Dempster Hwy, 18–20.vi.1980, 950 m, D.M. Wood & D.
Lafontaine (1 ♂, CNC); Richardson Mts, 2800 ft, 66°07′N 136°30′W, 11.vi.1982, D.M. Wood (1 ♂, 1 ♀, CNC).
RUSSIA. Chukotka AO (Wrangel Island): SE part of Wrangel Island, Rogers Bay, tundra, 20.vii.1932, A. Mineev
(2 ♂, ZIN); same data as holotype, 19.vii.1966, KBG (1 ♂, 1 ♀, ZIN; 1 ♂, 1 ♀, CNC); environs of Somnitelnaya
Bay, south of Wrangel Island, valley of river Somnitelnaya, Mineev Mtns, 150 m, shingle, on flower of Dryas
integrifolia, 19.vii.1966, KBG (2 ♀, ZIN); same locality, on flower of Potentilla emarginata, 22.vii.1966, KBG (1
♀, ZIN); Somnitelnaya Bay, south of Wrangel Island, Mineev Mtns, 400 m, S of Sovetskaya Mtn, 20.vii.1966, KBG
(2 ♀, ZIN); 5 km N Somnitelnaya Bay, warmed up on stones river bank, 27.vii.1966, KBG (1 ♀, ZIN); 5 km N
Somnitelnaya Bay, valley of river Somnitelnaya, small meadow, 26.vii.1972, KBG (1 ♀, ZIN); environs of Aterton
Mtn, 70°59′N 178°41′W, BT B, Sw, 13.vii.2019, OAK (1 ♂, 1 ♀, ZIN).
Additional material. RUSSIA, Chukotka AO (Wrangel Island): middle flow of river Neozhidannaya,
71°01′N 179°09′E, YPT, BT 1, 12–16.vii.2016, LFV (10 ♂, 10 ♀, ZIN; 1 ♂, 1 ♀, CULSP); same locality, YPT, BT
2, 12–16.vii.2016, LFV (37 ♂, 28 ♀ ZIN); Thomas Mtn, 70°56.964′N 178°43.026′E, BT 16, 10–31.vii.2016, LFV
(2 ♂, ZIN).
Diagnosis. Mid-sized (wing length about 4.5 mm), robust, black setose, blackish brown flies. Male holoptic;
hind tibia thickened, hind basitarsus strongly expanded, basal area broader than apex of hind tibia, elongate oval,
somewhat narrowed apically, both clothed in very long, dense fine setae; abdominal tergite 8 without projections;
cercus truncate at apex (lateral view), with notch slightly beyond middle dorsally; phallus well exposed, slender,
gently sinuate near middle. Female wing darker near base, legs without pennate setae.
Description. Body length 4.2–4.7 (holotype 4.7); wing length 4.3–4.5 (holotype 4.4) mm. Male (Fig. 62). Head
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 55
with dense greyish brown pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Holoptic;
upper ommatidia enlarged. Frons represented by very small subtriangular space below ocellar tubercle and larger
subtriangular space above antennae, bare. Face broad, bare. Ocellar triangle with several long fine setae. Occiput
with numerous long fine setae, including postoculars; postgena with numerous hair-like setae. Antenna blackish
brown; scape and pedicel subequally short, both with long fine setae on outer side and short setulae on inner side;
postpedicel conical, with straight margins, nearly 3X longer than wide; stylus very short, shorter than postpedicel
basal width. Palpus dark; with numerous long, dark fine setae. Proboscis with labrum dark reddish-brown, slightly
longer than head height.
Thorax dark in ground-colour, greyish brown pruinescent; with black setation; scutum almost uniformly
brownish pruinescent (dorsal view), only some traces of two narrow vittae between rows of acr and dc. Proepisternum
with tuft of numerous long fine setae on lower section; upper proepisternum in front of spiracle with numerous
similar setae. Prosternum bare. Antepronotum with numerous short setae. Postpronotal lobe with numerous long
fine setae. Mesonotal setae hardly differentiated, numerous, long, almost uniformly fine, only some npl, pal, sctl
and prescutellar dorsocentrals somewhat stronger; 1–2 pal, 6–10 sctl (usually 6–8); acr long, fine, arranged in 2
irregular rows, lacking on prescutellar depression; presutural dc long, multiserial, not separated by bare space from
supra-alar setae, some prescutellars somewhat stronger. Laterotergite with numerous long fine setae. Anterior and
posterior spiracles brown.
Legs robust, uniformly dark brown; mostly subshiny, coxae and trochanters with denser greyish pruinescence;
black setose. Fore coxa covered with long fine setae anteriorly. Podomeres of fore and mid legs unmodified; hind
femur somewhat broader near apical half; hind tibia expanded, slightly arcuate; hind basitarsus strongly expanded,
basal area broader than apex of hind tibia, elongate oval, somewhat narrowed apically; hind tarsomeres 2–3 slightly
expanded. Mid and hind femora whitish pilose ventrally. Legs mostly covered with fine setae, only mid and hind
tibiae with some stronger setae. Fore femur clothed in long setae including anteroventral and posteroventral rows
(about as long as femur width); mid femur with row of short anteroventrals and row of short posteroventral setae
on basal half and longer on apical half, some short setae near extreme base ventrally; hind femur without setae
ventrally, with several long av setae on about apical third (otherwise av minute), with more or less distinct row of
long setae anteriorly and some long setae around apex and base. Fore tibia with 2–3 more or less distinct ad (except
circlet of short subapicals), covered with moderately long fine setae posterodorsally; mid tibia with 2–3 ad, fine
setae posterodorsally (sometimes 1–2 setae somewhat stronger), rows of numerous short spine-like av and pv setae;
subapical circlet with 1 long ad seta; hind tibia clothed in very long dense curved setae; 1 seta in posteroapical comb.
Fore and mid basitarsi with some longer fine setae posterodorsally, spine-like setae ventrally; hind basitarsus with
numerous very long setae dorsally and ventrally (the latter stronger), hind tarsomeres 2–4 with long setae dorsally.
Wing membrane faintly brownish, with brownish veins; all veins complete (except Sc). Pterostigma brownish
yellow. Basal costal seta usually present, short (sometimes indistinct). Anal lobe well-developed; axillary incision
acute but closer to 90°. Squama brownish, dark fringed. Halter brown.
Abdomen dark, densely brownish pruinescent; covered with uniform black fine setae shorter on tergites dorsally
(except noted). Tergite 7 slightly convex, almost devoid of setation, with only 5–6 short posteromarginal setae
medially and some setulae laterally. Sternite 7 with similar structure and setation to sternite 6. Segment 8 with tergite
and sternite separated. Tergite 8 narrow, somewhat upturned posteriorly, slightly narrowly concave posteromedially;
with cluster of several long setae near posterior margin laterally. Sternite 8 large, somewhat convex on posterior
part ventrally, weaker sclerotized along midline (especially posteriorly); with numerous long setae along posterior
margin, otherwise bare.
Terminalia (Figs 63–65) brownish. Cerci separated from each other and from epandrium, not extended anteriorly
beyond tergite 8; mostly grey pruinescent, covered with dark mostly fine setulae, spinule-like setulae on inner face
subapically; cercus slightly shorter than epandrium, subrectangular and truncate at apex (lateral view), with notch
forming small projection slightly beyond middle dorsally; without pair of subepandrial lobes beneath. Epandrium
subtriangular, mostly grey pruinescent, subshiny anteriorly, with numerous long black setae along lower margin.
Hypandrium small, subtriangular (ventral view), narrowly sclerotized along margin; bare. Phallus (Fig. 65) well
exposed; mostly slender, evenly expanded on basal part, gently sinuate near middle, short apical part extended
beyond cerci. Ejaculatory apodeme not extended beyond basal curvature of phallus, with broad lateral wings and
narrow vertical wing.
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56 · Zootaxa 4848 (1) © 2020 Magnolia Press
FIGURES 62–65. Rhamphomyia sublongiseta sp. nov., male. 62. habitus, holotype, lateral view; 63. postabdomen, lateral
view; 64. terminalia; 65. phallus, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—epandrium;
hypd—hypandrium; ph—phallus.
Female. Similar to male, except frons broad, with marginal setulae; eye ommatidia equally small; ocellar
tubercle and occiput with shorter and stronger setae. Thorax with shorter setation; mesonotum with somewhat
sparser and more differentiated setae, 3–4 npl, 6–8 sctl. Legs without pennate setae, shorter setose; mid femur with
dense uniformly long posteroventral setae, mid tibia without av and pv spine-like setae; hind femur less distinctly
narrowed on basal part, with longer av setae on basal part, dense long posteroventral setae; hind tibia and basitarsus
only slightly expanded (latter as broad as hind tibia at apex), with rather short setae dorsally and ventrally (nearly
as long as hind tibia or basitarsus width); tarsomeres 2–4 rather slender, with short setae. Wing membrane darker
near wing base (about level of cells br and bm); basal costa seta usually distinct, short. Abdomen densely light grey
pruinescent; cercus long slender, with dark setulae.
Distribution. Holarctic; in Eurasia R. sublongiseta sp. nov. is recorded only from Wrangel Island; in North
America this species is known only from Yukon.
Etymology. The specific name refers to the similarity of the new species to R. longiseta Saigusa, 1964.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 57
Remarks. The new species belongs to the R. hirtula group (Saigusa 1964; Sinclair et al. 2019). In the structure
and setation of the male hind leg, R. sublongiseta sp. nov. resembles R. longiseta known from the alpine region of
Hokkaido, Japan (Saigusa 1964). However, the new species differs from R. longiseta by the more extensively setose
hind tibia (with long setae only dorsally in R. longiseta), simple abdominal tergite 8 (with pair of short spatulate
projections in R. longiseta), brownish pruinescent abdomen (ash grey in R. longiseta) and darker wing base of the
female.
Habitat. On Wrangel Island this species was collected mostly in the colder regions (Neozhidannaya River and
some coastal areas), where it was captured in various dry biotopes. Apparently, the species is characterized by sharp
interseasonal fluctuations in numbers, as evidenced by both Gorodkov’s data (who collected it only in 1966) and
ours (on the Neozhidannaya River this species was numerous in 2016, but was not recorded in 2019).
Rhamphomyia taimyrensis Frey
(Figs 66, 67)
Rhamphomyia (Dasyrhamphomyia) taimyrensis Frey, 1950: 109. Type localities: Krasnoyarskiy Territory, shore of Taymyr
Lake near mouth of Siruta-Yamu River; Amu-Tarida River, north-western Taymyr, Russia.
Other references: Frey, 1955b: 520 (revision).
Rhamphomyia (Rhamphomyia) taimyrensis Frey: Chvála & Wagner, 1989: 291 (catalogue); Shamshev, 2001b: 342 (key); Yang
et al., 2007: 189 (catalogue); Shamshev, 2016: 82 (checklist).
Material examined. RUSSIA. Arkhangelskaya Prov. (Novaya Zemlya Archipelago): Chekin Bay, 1.vii.1901,
Timofeev (1 ♂, 1 ♀, ZIN); Matochkin Shar, radio station, 8.vii.1924, Tolmachow (1 ♀, ZIN). Chukotka AO
(Wrangel Island): Somnitelnaya Bay, on flowers of Papaver, 18.vii.1966, KBG (1 ♀, ZIN); same locality, on
flowers of Dryas integrifolia, spotted tundra, 19.vii.1966, KBG (8 ♂, 11 ♀, ZIN; 1 ♂, 1 ♀, CNC); same locality,
sea shore at midnight on flowers of Papaver polaris, 25.vii.1966, KBG (5 ♂, 3 ♀, ZIN); same locality, on flowers
of Potentilla emarginata, 26.vii.1966, KBG (3 ♀, ZIN); same locality, spotted tundra near village, 9.vii.1972, KBG
(5 ♂, 7 ♀, ZIN); 5 km N Somnitelnaya Bay, 28.vii.1971, KBG (1 ♂, 1 ♀, ZIN); Somnitelnaya Bay, valley of river
Somnitelnaya, southern slope of Mineev Mtn, 100 m, shingle, 26.vii.1966, KBG (3 ♀, ZIN); environs of Somnitelnaya
Bay, valley of river Somnitelnaya, Mineev Mtns, 150 m, shingle, on flower of Dryas integrifolia, 22.vii.1966, KBG
(1 ♂, ZIN); Somnitelnaya River, 71°00′N 179°32′W, BT 9ВG, 14.vii.2006, OAK (1 ♂, 1 ♀, ZIN); same locality
and biotope, 18.vii.2015, OAK (1 ♀, ZIN) environs of Tundrovaya Mtn, near stream, 18.vii.1972, KBG (1 ♂, 2 ♀,
ZIN); middle flow of Mamontovaya River, N Perkatkun, osier-bed in valley of river, 17.vii.1972, KBG (2 ♂, 1 ♀,
ZIN); same locality, 71°10′N 179°45′W, Arctic fox burrow, 11.vii.2006, OAK (2 ♂, 2 ♀, ZIN); same locality and
biotope, 11–21.vii.2006, OAK (1 ♂, ZIN); same locality and biotope, 1.vii.2015, OAK (1 ♂, ZIN); same locality,
BT 6, 9.vii.2006, OAK (2 ♂, 1 ♀, ZIN); same locality, BT 5А, 20.vii.2006, OAK (3 ♂, 15 ♀, ZIN); same locality,
BT 5, 30.vi.2015, OAK (3 ♂, 10 ♀, ZIN); same locality, BT 10, 8.vii.2006, OAK (4 ♂, 3 ♀, ZIN); same locality and
biotope, 30.vi.2015, OAK (2 ♂, ZIN); same locality, BT 17, 3.vii.2015, OAK (2 ♂, 1 ♀, ZIN); same locality, BT
1А, YPT, 2–4.vii.2015, OAK (1 ♂, ZIN); same locality, house, 30.vi.2015, OAK (1 ♀, ZIN); spurs of Pervaya Mtn,
71°09′N 179°27′W, BT 13, 28.vi.2015, OAK (2 ♂, 8 ♀, ZIN); same locality and biotope, 28.vi.–13.vii.2015, OAK
(2 ♀, ZIN); same locality, BT 15, 28.vi.2015, OAK (1 ♂, 1 ♀, ZIN); same locality and biotope, 28.vi.–13.vii.2015,
OAK (1 ♀, ZIN); upper flow of Neizvestnaya River, 71°13′N 179°19′W, BT C4, 3.vii.2006, OAK (11 ♀, ZIN);
same locality and biotope, 12–29.vii.2015, OAK (1 ♀, ZIN); same locality, BT С2, 21.vi.2015, OAK (2 ♀, ZIN);
same locality, BT С3, 21.vi.2015, OAK (2 ♀, ZIN); same locality, BT 10А, 22.vi.2015, OAK (1 ♂, ZIN).
Diagnosis. Mid-sized (wing length 4.5–5.3 mm) blackish flies. Male holoptic; body black setose; scutum greyish
brown pruinescent, without vittae; acr biserial, presutural dc 3–4-serial; hind tibia curved inward closer to base, hind
basitarsus expanded, nearly as broad as apex of hind tibia; halter with yellow knob; CuA+CuP complete; phallus
well exposed, long, with narrow U-shaped loop just beyond cerci. Female mid and hind femora with anterodorsal
and posteroventral rows of pennate setae (except extreme base); hind tibia with some subpennate setae dorsally and
ventrally.
Redescription. Body length 4.6–5.5; wing length 4.5–5.3 mm. Male (Fig. 66, described for first time). Head
with greyish brown pruinescence on face, frons, postgena, ocellar triangle and occiput; dark setose. Holoptic;
upper ommatidia enlarged. Frons represented by very small subtriangular space below ocellar tubercle and larger
SHAMSHEV ET AL.
58 · Zootaxa 4848 (1) © 2020 Magnolia Press
subtriangular space above antennae, with scattered marginal setulae; with single setulae or bare. Face broad,
bare. Ocellar triangle with 2 long and several short fine setae. Occiput with numerous long thin setae, including
postoculars; postgena with numerous finer setae than occiput. Antenna blackish brown; scape and pedicel short,
scape slightly longer, both with short setulae; postpedicel conical, slightly concave on apical part ventrally, nearly
2.5–3X longer than wide; stylus rather long, 1.3–1.5X shorter than postpedicel. Palpus dark; with long, dark fine
setae. Proboscis with labrum dark reddish-brown, 1.3–1.5X head height.
Thorax dark in ground-colour, greyish brown pruinescent; with black setation; scutum greyish brown
pruinescent, somewhat lustrous, without vittae. Proepisternum with tuft of numerous long setae on lower section;
upper proepisternum in front of spiracle without setae. Prosternum bare. Antepronotum with numerous short setae.
Postpronotal lobe usually with 2 long, stronger and several short fine setae (sometimes with additional long fine
setae). Mesonotal setation (number, robustness and position of main mesonotal setae very variable, often even on
right and left sides): several long fine setae on supra-alar space (psut spal more numerous), 3–5 npl (additional fine
setae anteriorly), 1 long and 1 short pal, 6–8 sctl (often with additional short fine setae); acr long, thin, arranged in
2 irregular rows, absent on prescutellar depression; presutural dc long, arranged in 3–4 irregular rows, postsuturals
1–2-serial, 3–4 prescutellars longest. Laterotergite with numerous long fine setae. Anterior and posterior spiracles
brown.
Legs uniformly dark brown; mostly subshiny, coxae and trochanters with denser greyish pruinescence; black
setose. Fore coxa covered with long fine setae anteriorly. Femora and tibiae slender; hind femur only slightly thicker
than fore and mid femora; hind tibia curved inward closer to base (posterior view); hind basitarsus thickened,
nearly as broad as hind tibia at apex. Mid and hind femora whitish pilose ventrally; fore femur with rows of fine,
long anteroventral and posteroventral setae (nearly as long as femur width) and similar setae anteriorly; mid femur
with rows of short fine anteroventral and long posteroventral setae; hind femur with complete row of rather long
fine anteroventral setae. Fore tibia with 2–3 short anterodorsal and long fine setae posterodorsally; mid tibia with
3–4 anterodorsal and 3–4 posterodorsal setae (except circlet of subapicals) becoming longer toward apex of tibia
(longest setae nearly 1.5X longer than tibia width), several short spine-like anteroventral and posteroventral setae on
about apical half. Hind tibia with numerous long anteroventral and posteroventral setae, lacking setulae on concave
space near base; 1 seta in posteroapical comb of hind tibia. Fore basitarsus with somewhat longer setae dorsally
(nearly as long as basitarsus width); mid and hind basitarsi with numerous long setae dorsally (longest setae nearly
twice longer that corresponding basitarsus width).
Wing membrane faintly brownish, mostly with brownish veins; all veins complete (except Sc); CuA+CuP (anal
vein) weakened on subapical portion. Pterostigma brownish yellow; 1 basal costa seta present, short. Anal lobe well-
developed; axillary incision almost 90°. Squama brownish, dark fringed. Halter with yellow knob and brownish
yellow stem.
Abdomen dark brown, faintly brownish pruinescent, subshiny; covered with uniform black fine setae shorter
on tergites dorsally, sternite 8 with numerous very long posteromarginal setae. Segments 6–7 with unmodified
vestiture and structure. Segment 8 with tergite and sternite separated, both normally sclerotized; tergite 8 concave
posteromedially; sternite 8 unmodified, subrectangular viewed laterally, nearly as long as sternite 7.
Terminalia dark brown (Fig. 67). Cerci directed posteriorly, separated from each other and from epandrium;
covered with numerous fine dark setulae dorsally, numerous spinule-like setae along lower margin; cercus
subrectangular (lateral view), with tapered apex, about as long as epandrium and half as broad as epandrium;
without pair of subepandrial lobes beneath. Epandrium subrectangular, broadly rounded apically; shiny on basal
region; covered with long, fine dark setae. Hypandrium mostly membraneous, only lateral arms sclerotized, bare.
Phallus yellowish to brownish yellow, well exposed; evenly curved and uniformly slightly broader on about basal
half until lower margin of epandrium, tapered apically; slightly curved between cerci, with narrow U-shaped loop
just beyond cerci. Ejaculatory apodeme not extended beyond basal curvature of phallus, with narrow lateral wings
and without prominent vertical wing.
Female. Similar to male, except face broad with lateral setulae; mid and hind femora with anterodorsal and
posteroventral rows of pennate setae (except extreme base); hind tibia with some subpennate setae dorsally and
ventrally. Cercus long slender, covered with dark setulae.
Remarks. The male was associated with the female after a pair taken in copula, which was found among
material collected from the type locality. The female was compared with the syntypes housed in ZIN.
Frey (1950) placed R. taimyrensis in his subgenus Dasyrhamphomyia, but this is not confirmed by our discovery
of a male of this species. Actually, R. taimyrensis is most closely related to R. laevigata Loew, revised very recently
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 59
by Sinclair et al. (2019), which is widely distributed in North America and also was known for a long time from the
extreme north of Russia as R. fridolini Frey. The subgeneric position of these species remains unclear.
Distribution. Palaearctic; R. taimyrensis is broadly distributed across the Russian Arctic including continental
areas and islands (Novaya Zemlya Archipelago, Wrangel).
FIGURES 66–68. Rhamphomyia. 66, 67. R. taimyrensis Frey. 66. male habitus, lateral view; 67. male terminalia, lateral view.
68. Rhamphomyia sp. 1, female habitus, lateral view. Abbreviations: cerc—cercus; ej apod—ejaculatory apodeme; epand—
epandrium; hypd—hypandrium; ph—phallus.
SHAMSHEV ET AL.
60 · Zootaxa 4848 (1) © 2020 Magnolia Press
Habitat. On Wrangel Island, this species was not found in the colder region (Neozhidannaya River), but in one
year (1966) it was numerous in the coastal spotted tundra. In the 2000s, this species was collected mainly in the
warmer central region of the island, in dry habitats, especially often in tundra-steppe-like vegetation.
Unnamed species of Rhamphomyia
Rhamphomyia sp. 1
(Fig. 68)
Material examined. RUSSIA. Chukotka AO (Wrangel Island): env. of Somnitelnaya Bay, south of Wrangel
Island, 10.vii.1966, KBG; upper flow of river Somnitelnaya, Mineev Mtns; on flower of Potentilla (1 ♀, ZIN).
Description. Female (Fig. 68). Length body 3.6 mm, wing 3.7 mm. Head with greyish brown pruinescence on
face, frons, postgena, ocellar triangle and occiput; dark setose. Eyes dichoptic, separated by broad frons. Antenna
blackish brown; scape and pedicel short, scape slightly longer, both with short setulae; postpedicel conical, 3X
longer than wide; stylus very short, nearly as long as postpedicel basal width. Palpus dark; with several moderately
long, dark fine setae. Labrum short, nearly of head height. Thorax brownish grey pruinescent; with black setation;
scutum almost uniformly faintly brownish grey pruinescent, without vittae in dorsal view, with some traces of darker
narrow vittae between acr and dc rows viewed anterodorsally. Proepisternum with few moderately long fine setae on
lower section; upper proepisternum in front of spiracle without setae. Prosternum bare. Antepronotum with several
short setae. Postpronotal lobe with 2 long and several shorter fine setae. Mesonotal setae fine, hardly differentiated;
2 longer presut spal with numerous additional short setae, 3–4 npl (with additional short setae), 1 psut spal (with
several additional short setae anteriorly), 1–2 pal, 6 sctl; acr arranged in 2 irregular rows, lacking on prescutellar
depression, shorter than presutural dc; presutural dc 3–4-serial, long, barely separated from numerous supra-alars,
postsutural dc 1–2-serial, longer, 3–4 prescutellars longest. Laterotergite with numerous long fine setae. Anterior
and posterior spiracles brown. Legs long, slender, uniformly brownish, faintly greyish pruinescent, dark setose; fore
coxa with fine setae anteriorly; femora and tibiae covered with short setae, no strong and pennate setae; hind femur
with very short anteroventral setae; tibiae with some longer setae dorsally, hind tibia with 1 seta in posteroapical
comb. Wing membrane uniformly brownish infuscate; CuA+CuP (anal vein) incomplete; 1 short basal costal seta;
axillary incision right-angled. Squama brownish, dark fringed. Halter with brownish knob and reddish brown stem.
Abdomen brownish, faintly greyish pruinescent, with short fine dark setae; cercus long, slender, with dark setulae.
Remarks. This species may belong to the R. pusilla species group of Pararhamphomyia, but this requires
verification. According to Frey, this species would run to his group III.
Family HYBOTIDAE
Subfamily Tachydromiinae
Genus Platypalpus Macquart
Platypalpus septentrionalis Shamshev & Sinclair sp. nov.
(Figs 69–72)
urn:lsid:zoobank.org:act:67F5F184-B4CB-4F4E-895D-491D576A1A5E
Type material. HOLOTYPE ♂, labelled: [RUSSIA. Chukotka AO (Wrangel Island)] “bukh. [=bukhta,
bay] Somnitelnaya,/ south o. [=ostrov, island] Wrangel/ Gorodkov 22.vii.[1966]”; “dol. r. [=dolina reki, valley of
river] Somnitelnaya,/ Mineev Mountains, 150 m,/ shingle”; “on flower of Potentilla emarginata”; “Holotypus/
Platypalpus/ septentrionalis/ Shamshev, Sinclair sp. nov. [red label]” (INS_DIP_0000619, ZIN, pinned on one pin
with a female).
PARATYPES. RUSSIA. Chukotka AO (Wrangel Island): Pinned on one pin with the holotype male (1 ♀,
ZIN); same data as holotype (1 ♂, 4 ♀, ZIN; 1 ♂, 1 ♀, CNC); same data as holotype, 19.vii.1966 (1 ♀, ZIN).
Diagnosis. Small robust blackish species of the P. longicornis group; 2 pairs of black vertical setae; postpedicel
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 61
nearly 2.5X longer than wide and 2X longer than stylus; legs extensively dark brown, mid tibia with very short
apical spur; halteres brown.
Description. Body holotype 1.9 mm, wing 2.0 mm. Male (Fig. 69). Head black. Occiput entirely densely
greyish pollinose; with 2 pairs of long, black vertical setae (inner pair proclinate and slightly inclinate; outer pair
lateroclinate); very short dark setae on upper part and longer pale setae on lower part. Ocellar tubercle concolorous
with occiput, with 2 long, black lateroclinate anterior setae and 2 minute posterior setae. Frons broad on upper
part, narrowed toward antennae, below ocellar tubercle broader than distance between outer margins of posterior
ocelli; densely greyish brown pruinose (anterior view). Face rather broad, nearly as broad as frons above antennae,
concolorous with frons; clypeus shiny. Antenna uniformly dark brown; postpedicel conical, moderately long, nearly
2.5X as long as basal width; stylus short, nearly 2X shorter than postpedicel. Gena very narrow, shiny. Proboscis
dark brown, short. Palpus black, small, oval, pale pubescent, with 2 pale setae apically.
Thorax black, almost entirely uniformly greyish tomentose; katepisternum almost entirely shiny, only narrowly
tomentose along upper margin and anteriorly. Postpronotal lobe with 1 long black seta and some minute setulae.
Mesonotum with main setae black to brown; 2 long npl (with additional pale setulae), 2 pal (1 much longer), 4 sctl
(apical pair long, lateral pair short); acr very short, fine, arranged in 2 broad irregular rows; dc mostly arranged in 2
irregular rows and almost similar to acr anteriorly (some setae very slightly longer), 2 prescutellar pairs long.
Legs extensively dark brown, subshiny; fore femur narrowly yellowish brown apically, mid and hind femora
reddish brown on extreme apex; fore tibia mostly yellowish brown, somewhat paler closer to base, mid and hind
tibiae reddish brown near extreme base. Coxae broadly shiny laterally, otherwise faintly greyish pruinose; fore coxa
with pale setae anteriorly; hind trochanter with fine setulae. Fore femur strongly thickened; pilose ventrally; with
anteroventral and posteroventral mostly minute fine setae becoming slightly longer toward base. Mid femur slightly
stouter than fore femur; with double row of blackish short spine-like setae ventrally, without row of posteroventral
setae. Hind femur slender, with minute anteroventral setae. Fore tibia slightly uniformly thickened; with simple
setulae dorsally. Mid tibia slightly arcuate; with row of ventral dark spinule-like setulae; apical spur very short,
shorter than apical width of tibia. Hind tibia slender, straight, with simple setulae. Tarsi of all legs unmodified.
Wing faintly infuscate, with pale brownish veins. Basal costal seta long, blackish. Veins R4+5 and M1+2 parallel
near wing margin; CuA slightly recurrent; anal vein very faint; crossveins m-cu and r-m somewhat contiguous.
Calypter brown, pale fringed. Halter brown.
Abdomen entirely dark brown to black, almost entirely shiny, only tergites 1 and 2 laterally faintly greyish
pruinose (tergite 2 only narrowly anteriorly); covered with scattered pale setulae.
Terminalia (Figs 70–72) small, subglobular, concolourous with abdomen, epandrium mostly shiny. Right cercus
(Fig. 71) digitiform with somewhat pointed apex, short, with small internal projection slightly beyond middle;
covered with short simple setae. Left cercus of similar shape to right cercus viewed dorsally, broadly ovate viewed
laterally, nearly as long and as broad as right cercus (dorsal view); with small internal projection slightly beyond
middle; covered with short simple setae. Hypoproct subtriangular (dorsal view), with somewhat pointed apex. Right
epandrial lamella (Fig. 70) subglobular viewed laterally, with row of several short to moderately long subapical
setae; right surstylus hardly differentiated from apex of epandrial lamella, short, rounded apically. Left epandrial
lamella (Fig. 72) rather subtriangular but with small rounded projection about mid-dorsally; with scattered short
setae along ventral margin and some setulae ventrally.
Female. Similar to male. Visible parts of tergites 7 and 8 densely greyish pruinescent; cercus long, slender.
Distribution. Palaearctic: Russia (Chukotka Autonomous Okrug, Wrangel Island).
Etymology. The specific name is from the Latin septentrionalis (north, northern), in reference to the distribution
of this species.
Remarks. This new species can be readily distinguished from all other species of Holarctic Platypalpus
by brown halteres. In having small body size, two pairs of vertical setae, very short midtibial spur and lacking
posteroventral setae on the mid femur, the new species resembles P. vegrandis Frey, 1943 known from Europe
(Chvála 1989). However, P. vegrandis has partly yellow antennae and legs. In addition, hind trochanters in the male
of this species bear three black spine-like setae. It is interesting to note that both species are found on flowers and
may have similar hunting behaviour (Grootaert et al. 2012).
Also, the new species could be compared with P. palmeni Frey, 1943 known from the Alps (Chvála 1989). Both
species are similar in body size, have two pairs of vertical setae and lack posteroventral setae on the mid femur.
However, P. palmeni can be readily distinguished from the new species by pale yellow palps and extensively yellow
legs.
SHAMSHEV ET AL.
62 · Zootaxa 4848 (1) © 2020 Magnolia Press
FIGURES 69–72. Platypalpus septentrionalis sp. nov., male. 69. habitus, holotype, lateral view; 70. right epandrial lamella
and right cercus, lateral view; 71. epandrium with cerci, dorsal view; 72. left epandrial lamella and left cercus, lateral view.
Abbreviations: cerc—cercus; epand—epandrium; hypd—hypandrium; rt sur—right surstylus.
In addition, the new species was compared with the syntypes (currently in CNC) of the following species
known from subarctic areas of North America in order to confirm its status (especially since the Nearctic fauna
is not as well diagnosed in comparison to the Palaearctic): Platypalpus arcticus Melander, 1928, P. diversipes
Coquillett, 1900, P. gilvipes Coquillett, 1900, P. lateralis Loew, 1864, P. vittiger Melander, 1928 (described as var.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 63
of P. flavirostris Loew, 1864). All these species differ from the new species primarily by a combination of characters
noted in the diagnosis.
Platypalpus subtectifrons Shamshev & Sinclair sp. nov.
(Figs 73–76)
urn:lsid:zoobank.org:act:FC7E4A07-9126-4941-A797-489575CA0E21
Type material. HOLOTYPE ♂, labelled: [RUSSIA. Chukotka AO (Wrangel Island)] “sr. tech. r. [=srednee
techenie reki, =middle flow of river] Mamontovaya, N/ Perkatkun, o. [=ostrov, island] Wrangel/ willows in river
valley/ Gorodkov 17.vii.[1]972”; “Holotypus/ Platypalpus/ subtectifrons/ Shamshev, Sinclair sp. n. [red label]”
(INS_DIP_0000620, ZIN).
PARATYPES. CANADA. Northwest Territories: Aklavik, 2.vii.1956, R.E. Leech (1 ♀, CNC); same data
except, 25.vi.–3.vii.1956, E.F. Cashman (3 ♀, CNC); Fort McPherson, 17–19.vii.1957, R. Hurley (2 ♀, CNC);
21 mi E Tuktoyaktuk, 8–12.vii.1972, D.M. Wood (2 ♂, 1 ♀, CNC); Tuktoyaktuk, 69°26′16.2″N 133°1′2.16″W,
pan traps, meadow, 15–18.vii.2010, Goulet & Boudreault (12 ♂, 4 ♀, CNC). Yukon: British Mtns, Firth River,
25.vii.1956, E.F. Cashman (2 ♀, CNC); Firth River, 14.vii.–7.viii.1956, E.F. Cashman (5 ♂, 7 ♀, CNC); same data
except, 20.vii.–3.viii.1965, R.E. Leech (5 ♂, 8 ♀, CNC). RUSSIA. Chukotka AO (Wrangel Island): Same data
as holotype (4 ♂, 1 ♀, ZIN; 2 ♂, 1 ♀, CNC); 21.vii.1972, KBG (1 ♂, 4 ♀, ZIN); middle flow of river Neizvestnaya,
71°20.287′N 179°29.779′W, BT 1, 5–14.vii.2018, U.V. Babiy (3 ♂, 3 ♀, ZIN); middle flow of river Mamontovaya,
71°10′N 179°45′W, BT 2, 30.vi.–9.vii.2006, OAK (1 ♂, ZIN); same locality and biotope, BT 2, YPT, 2–4.vii.2015,
OAK (2 ♂, ZIN); same locality, 71°10′N 179°45′W, BT 1A, YPT, 2–4.vii.2015, OAK (3 ♂, 3 ♀, ZIN); same locality,
71°10′N 179°45′W, BT 14, Sw, 2.vii.2015, OAK (2 ♂, 1 ♀, ZIN); same locality, 71°10′N 179°45′W, 3.vii.2015,
OAK (1 ♂, ZIN); Tundrovaya Mnt, 71°18′N 179°49′W, BT 2, 1–19.vii.2015, LFV (1 ♂, ZIN; dissected). USA.
Alaska: Cape Thompson, 23–25.vii.1961, R. Madge (2 ♀, CNC); Isabel Pass, mi 206 Richardson Hwy, 2900 ft,
18.vii.1962, P.J. Skitsko (2 ♀, CNC); Nome Area, mi 46 Kougarok Rd, 64°57′27″N 168°48′21″W, 26–28.vi.2005,
MT, J. & R. Skevington (1 ♂, CNC); Prudhoe Bay, 9.vii.1971, M. Deyrup (1 ♂, 6 ♀, CNC); Umalakleet, 18,28.
vi.1961, B.S. Heming (1 ♂, 1 ♀, CNC); same data except, 28.vi., 8.vii.1961, R. Madge (1 ♂, 1 ♀, CNC); Umiat,
3–25.vii.1959, R. Madge (3 ♂, 38 ♀, CNC); same data except, 6–15.vii.1959, J.E.H. Martin (26 ♂, 28 ♀, CNC);
same data except, 4.viii.1959, B.N. Smallman (1 ♂, CNC).
Diagnosis. Small, blackish species of the P. pallidiventris group; one pair of vertical setae, postpedicel nearly
1.5X longer than wide, palpus black, thorax with pale setae, scutum entirely pruinose, acr biserial, legs almost
entirely dark brown, hind tibia with long apical spur, abdomen almost entirely shiny.
Description. Body length 2.4–2.5 (holotype 2.5), wing length 3.0–3.2 (holotype 3.1) mm. Male (Fig. 73). Head
black, mostly pale setose, ocellar setae rather yellowish. Occiput entirely densely greyish pruinose; with one pair of
moderately long, fine, wide apart vertical setae; numerous short setae on upper part and longer setae on lower part.
Ocellar tubercle concolorous with occiput, with 2 long, lateroclinate anterior setae and 2 minute posterior setae.
Frons broad, only slightly narrowed toward antennae, below ocellar tubercle broader than distance between outer
margins of posterior ocelli; densely light grey pruinose (anterior view). Face broad, almost as broad as frons above
antennae, densely whitish pruinose (including clypeus). Antenna uniformly dark brown; postpedicel short, nearly
1.5X (in holotype 1.65X) as long as basal width; stylus nearly 2X (in holotype 2.2X) longer than postpedicel. Gena
narrow, shiny. Proboscis dark brown, elongate, nearly 1.5X shorter of head height. Palpus black, small, oval, whitish
pubescent, with few short setae.
Thorax black, mostly pale setose, strong setae rather pale yellowish; prothoracic sclerites and mesopleuron
densely greyish tomentose, scutum rather yellowish grey faintly tomentose; katepisternum mostly shiny, only
tomentose on upper anterior corner and along posterior corner. Postpronotal lobe with 1 moderately long fine seta
and some minute setulae. Mesonotum with 2 moderately long npl (with numerous additional setulae), 2 pal (1
longer), 4 sctl (apical pair long, lateral pair finer and shorter; sometimes additional setae present); some setulae
behind postpronotal lobe and on supra-alar space; acr short, fine, arranged in 2 moderately broad irregular rows, on
prescutellar depression hardly distinguishable from dc and sometimes more numerous; dc arranged in 2–3 irregular
rows and mostly similar to acr, some prescutellars longer.
Legs extensively dark brown, subshiny, pale setose (except noted), somewhat variable in yellowish pattern. In
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64 · Zootaxa 4848 (1) © 2020 Magnolia Press
holotype and most paratypes fore femur very narrowly yellowish apically, mid and hind femora yellowish brown
on extreme apex, fore tibia yellowish near base (becoming brownish toward apex); mid and hind tibiae slightly
paler than corresponding femora, tarsomeres somewhat paler on basal part (broadly on basitarsus, extreme base
on remaining tarsomeres); in addition, in some paratypes fore coxa on extreme apex and all femora on extreme
base yellowish brown (1 ♂ and 5 ♀ from Perkatkun). Fore coxae broadly shiny laterally, otherwise densely greyish
tomentose; mid and hind coxae almost entirely shiny; all coxae pale setose; hind trochanter with fine setulae. Fore
femur strongly thickened; pilose ventrally; with rows of long, fine anteroventral and posteroventral setae (longest
setae on basal part slightly longer than half of femur width). Mid femur slightly stouter than fore femur; with double
row of short, blackish spine-like setae ventrally, row of long strong posteroventrals intermixed with shorter and
finer setae. Hind femur slender, with somewhat longer anteroventral setae on about apical half. Fore tibia slightly
uniformly thickened; with simple setulae dorsally, some anterodorsals slightly stronger; longer setae ventrally and
posteroventrally (longest ones on subapical part slightly longer than tibia width). Mid tibia with row of ventral
dark spinule-like setulae; apical spur long, pointed, with two setulae at tip. Hind tibia slender, straight, with simple
setulae. Tarsi of all legs unmodified; fore basitarsus with longer setae ventrally and posteroventrally (similar to
those on fore tibia).
Wing faintly infuscate, with brownish veins. Basal costal seta moderately long, fine, pale. Veins R4+5 and M1+2
slightly convergent near wing margin; CuA straight, perpendicular, faint apically; anal vein very faint; crossveins
m-cu and r-m slightly separated. Calypter dirty yellow, pale fringed. Halter yellow.
Abdomen entirely blackish brown, almost entirely shiny, only tergite 1 faintly greyish pruinose laterally; covered
with numerous pale hair-like setae longer on tergites laterally.
Terminalia (Figs 74–76) small, concolourous with abdomen, epandrium shiny. Right cercus (Fig. 75) digitiform,
long, almost parallel-sided toward apex, truncate; covered with short simple setae. Left cercus digitiform, slightly
longer and narrower (on apical half) than right cercus (dorsal view), almost parallel-sided toward apex, rounded on
apex; covered with short simple setae. Hypoproct subtriangular, narrow (dorsal view), with pointed apex almost
reaching apex of cerci. Right epandrial lamella (Fig. 74) subtriangular viewed laterally, with row of several long
subapical setae; right surstylus well differentiated from apex of epandrial lamella, long, slender, with some minute
marginal setulae. Left epandrial lamella subtriangular (Fig. 76) somewhat concave about mid-ventrally; with
numerous long pale yellow setae along about apical 2/3 of ventral margin.
Female. Similar to male. Visible parts of tergites 7 and 8 densely greyish pruinescent (except narrow upper
margin of sternite 8); cercus long, slender.
Distribution. Palaearctic: Russia (Chukotka Autonomous Okrug, Wrangel Island). In North America, this
species is known from Alaska, Northwest Territories and Yukon.
Etymology. The specific name refers to the similarity of the new species to P. tectifrons (Becker).
Remarks. This new species strikingly resembles the poorly known P. tectifrons (Becker, 1907) described from
eastern Tibet of China (syntypes housed in ZIN), which has the same pattern of leg colour and other main characters.
The redescription of P. tectifrons will be published in a forthcoming paper. Platypalpus tectifrons differs from the
new species by the somewhat narrower, longer postpedicel (about 2.5X longer than wide) and brownish to brownish
yellow main mesonotal setae. Within the key to Platypalpus compiled by Chvála (1989), the new species would
run to P. interstinctus (Collin, 1926) and P. pseudofulvipes (Frey, 1909) (P. pallidiventris group). Platypalpus
interstinctus (somewhat variable in leg colour) differs from the new species by distinctly black annulated tarsi,
brownish vertical, acrostichal and dorsocentral setae, as well as by details of the male terminalia (Chvála 1975:
178, figs 441–443). Platypalpus pseudofulvipes can be readily distinguished from the new species primarily by the
spindle-shaped dilated fore tibiae. In addition, the new species could be compared with P. melancholicus (Collin,
1961), P. annulatus (Fallén, 1815) and P. carteri (Collin, 1926). Platypalpus melancholicus and P. annulatus differ
from the new species primarily by 4-serial acrostichals (at least on anterior part of mesoscutum). Platypalpus carteri
can be distinguished from the new species by lateral pruinosity on all abdominal tergites. The male terminalia of
the new species somewhat resemble those of P. notatus (Meigen, 1822) (especially cerci), however, P. notatus has
longer postpedicel (about 2.5X longer than wide) and entirely shiny abdomen (including lateral area of first two
tergites).
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 65
FIGURES 73–76. Platypalpus subtectifrons sp. nov., male. 73. habitus, holotype, lateral view; 74. right epandrial lamella,
lateral view; 75. epandrium with cerci, dorsal view; 76. left epandrial lamella and left cercus, lateral view. Abbreviations:
cerc—cercus; epand—epandrium; hypd—hypandrium; rt sur—right surstylus.
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66 · Zootaxa 4848 (1) © 2020 Magnolia Press
In addition, the new species was compared with the syntypes (currently in CNC) of the species known from
subarctic areas of North America listed under above. All these species differ from the new species primarily by a
combination of characters noted in the diagnosis.
Habitat. On Wrangel Island, this species was collected along riverbeds.
Genus Tachypeza Meigen
Tachypeza sericeipalpis Frey
(Figs 77–80)
Tachypeza sericeipalpis Frey in Lundström & Frey, 1913: 10. Type locality: Cape Mikulkin (Kanin Peninsula, 67°48′N 46°41′E),
Nenets Autonomous Okrug, Russia.
Other references: Frey, 1915: 14 (faunistic survey); Chvála, 1975: 224 (revision); Shamshev, 2016: 148 (checklist).
Tachypeza dilutata Frey, 1915: 14 (as var. of sericeipalpis Frey). Type locality: Kharaulakhskij Ridge near mouth of the Lena
River (~71°44′N 128°16′E), Yakutia, Russia.
Material examined. CANADA. Northwest Territories: Ford Lake, 63°11′N 107°19′W, 23.vi.1966, G.E. Shewell
(1 ♂, 1 ♀, CNC). Nunavut: Whatever Lake, 62°41′N 97°03′W, vii.1989, pitfall trap, J. Pilny (3 ♂, 1 ♀, CNC).
RUSSIA. Chukotka AO (Wrangel Island): N Somnitelnaya Bay, 27.vii.1966, KBG (1 ♀, ZIN); northern slope of
Berry Peak, 7 km ESE Sovetskaya Mtn, 350 m, on stones, 12.vii.1972, KBG (5 ♂, 6 ♀, ZIN).
Recognition. Small blackish flies (body 2.5–3 mm) (Fig. 72); palpus pale to pale yellow, whitish pubescent,
with short pale hair-like setae (sometimes 1 subapical seta brownish); prothoracic sclerites (except postpronotal
lobe) densely silvery grey tomentose, mesopleuron almost entirely shiny, 1 strong npl, 2 broadly set similar sctl, 1
(sometimes 2) strong prescutellar dc; legs almost uniformly brown; halter with yellow knob. Male: mid femur with
row of 6–7 closely set black short anteroventral spine-like setae on about basal 1/3 and 2 closely set, moderately
long, spine-like setae near extreme base; mid tibia with shallow, ventral subapical excision. Male terminalia (Figs
78–80) large, brown, epandrium shiny. Cerci (Fig. 79) separated, shifted to inner margin of right epandrial lamella;
right cercus digitiform, short, with apex slightly extended beyond apex of right epandrial lamella, covered with
short setae stronger on inner margin; left cercus short, broad, subrectangular; with some short setae. Hypoproct
weakly sclerotized, rather subtriangular, separated from both cerci, bare. Right epandrial lamella (Fig. 78) without
ventral lobe, produced apicoventrally; with cluster of several spine-like short marginal setae apically; right surstylus
separated from epandrial lamella, narrow, with two projections, outer projection very short, inner projection long,
stylet-like; right bacilliform sclerite with 2 closely set spine-like setae. Left epandrial lamella narrow; with large
apodeme; bearing some setulae apically; left surstylus (Fig. 80) large, rather subtriangular (lateral view) but with
additional narrow pointed projection dorsally; with short marginal setae apically. Hypandrium hump-backed.
Distribution. Holarctic; T. sericeipalpis has a disjunct distribution occurring in Eurasia across the arctic region
of Russia from Murmanskaya Province to Chukotka (including Wrangel Island) and is found locally in mountains
of Altay Republic and Buryatia (Shamshev, unpubl. data). In North America, this species is newly recorded from the
Northwest Territories and Nunavut of Canada.
Remarks. The male terminalia of this species are here illustrated for the first time.
Discussion
Three families of flies of the superfamily Empidoidea (exclusive of Dolichopodidae) were found in our study from
the Russian Arctic islands and Svalbard Archipelago—Brachystomatidae, Empididae and Hybotidae (Table 1).
The Brachystomatidae are a small group comprising about 200 described species worldwide (Sinclair 2017),
it is better represented in the Southern Hemisphere. Only the genus Heleodromia (subfamily Trichopezinae) was
recorded from the Arctic islands (one uncertain species from Novaya Zemlya and two species from Wrangel).
Heleodromia includes currently 26 species distributed exclusively in the Holarctic and mountains of the Oriental
Realm (Himalayas) (Sinclair et al. 2011).
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FIGURES 77–80. Tachypeza sericeipalpis Frey, male, lateral view. 77. habitus, lateral view; 78. right epandrial lamella, dorsal
view; 79. epandrium with cerci, dorsal view; 80. left surstylus, lateral view. Abbreviations: cerc—cercus; epand—epandrium;
hypd—hypandrium; rt sur—right surstylus.
The Empididae are one of the largest families of Empidoidea. The group has worldwide distribution comprising
about 3500 described species. Empidids dominate absolutely on the Arctic islands (32 species, 89%) among
Empidoidea including two genera of the subfamily Clinocerinae (Clinocera, Trichoclinocera) and two genera of
Empidinae (Empis, Rhamphomyia). Clinocera and Trichoclinocera are quite small groups (109 and 39 described
species, respectively), the former is represented by two species (one uncertain) from Novaya Zemlya and Wrangel,
whereas the latter is known by a single species from Wrangel only. Empis and Rhamphomyia are two mega-diverse
genera of Empididae, with about 800 and 600 described species worldwide, respectively (Yang et al. 2007).
However, many more new species are still waiting description, especially in Rhamphomyia (Barták & Kubík
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68 · Zootaxa 4848 (1) © 2020 Magnolia Press
2009; Saigusa 2012). Empis is more diverse in southern regions and only very few species penetrate to the Arctic
territories (Shamshev 2016). A single species recorded from Wrangel Island is the extreme north finding of the
genus. Rhamphomyia is primarily a Holarctic group where it is especially diverse in temperate and boreal regions.
Considering its great diversity, it is not surprising that Rhamphomyia dominates absolutely among Empididae on the
Arctic islands (29 species, 91%) and Empidoidea on the whole.
FIGURES 81–86. Landscapes in different regions of Wrangel Island. 81. middle course of Neozhidannaya River, zonal spotted
tundra; 82. middle course of Neozhidannaya River, river valley; 83. coastal Southern plain; 84. environs of Somnitelnaya Bay,
Mineev Mtns; 85. central part of Wrangel Island, middle flow of Mamontovaya River, oseir-bed in the river valley; 86. central
part of Wrangel Island, spurs of Pervaya Mtn, herb-dryad dry spotted tundra.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 69
TABLE. Distribution of Empididae, Hybotidae and Brachystomatidae among Russian Arctic islands and Svalbard Archipelago (doubtful identifications are included with a ques-
tion mark).
Taxon Svalbard Kildin Novaya Zemlya Kolguev Vaygach Dikson New Siberian Wrangel Kolyuchin Total species
Brachystomatidae 2
Heleodromia 2
H. sp.? X
irwini X
pullata X
Total Brachystomatidae / territory 0 0 1 0 0 0 0 2 0 –
Empididae 32
Clinocerinae 2
Clinocera 1
C. sp.? X
nivalis X X
Trichoclinocera 1
lapponica X
Empidinae 30
Empis 1
septentrionalis sp. nov. X
Rhamphomyia 29
albopilosa X
albosegmentata X X
armipes X
brusewitzii X X
erinacioides X X X
filicauda X
frigida X
gorodkovi sp. nov. X
hirtula X
hoeli X
hovgaardii X X X X
......continued on the next page
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70 · Zootaxa 4848 (1) © 2020 Magnolia Press
TABLE (Continued)
Taxon Svalbard Kildin Novaya Zemlya Kolguev Vaygach Dikson New Siberian Wrangel Kolyuchin Total species
hybotina X
kaninensis X
kjellmanii X X X
longestylata X
mallochi sp. nov. X
morio X
nigrita X
nordqvistii X
septentrionalis X
shewelli X
subfilicauda sp. nov. X
sublongiseta sp. nov. X
submacrura sp. nov. X
subtenuiterfilata sp. nov. X
taimyrensis X X
ursinella X
wrangeli sp. nov. X
sp. 1 X
Total Empididae / territory 1 3 10 1 1 3 2 22 1 –
Hybotidae 3
Platypalpus 2
septentrionalis sp. nov. X
subtectifrons sp. nov. X
Tachypeza 1
sericeipalpis X
Total Hybotidae / territory 0 0 0 0 0 0 0 3 0 –
Total species / territory 1 3 10 1 1 3 2 27 1 37
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 71
The Hybotidae are the second of the largest families of Empidoidea. The group has worldwide distribution
comprising about 2300 described species. The hybotids are especially diverse in tropical regions. However, some
broadly distributed genera of this family exhibit high levels of species diversity in temperate regions as well (e.g.,
Platypalpus). Although several genera of Hybotidae are primarily Holarctic in distribution, they are absent on the
Arctic islands. Three species belonging to two genera of the subfamily Tachydromiinae (Platypalpus—2 species,
Tachypeza—1 species) were found in our study from Wrangel Island only.
In addition, we should mention Dolichopodidae, which is the largest family of Empidoidea with about 8000
described species (Grichanov 2017). This huge cosmopolitan group appears to be very poorly represented on the
Arctic islands. Only six species have been recorded from the Russian territories (Kolguev—1, Dolgiy—1, Wrangel—
5) and there are no species from Svalbard, Novaya Zemlya, Severnaya Zemlya and New Siberian Archipelagos
(Grichanov 2006; Coulson 2007; Grichanov & Khruleva 2018). A similar situation is typical for the islands of the
Queen Elizabeth Islands (Canada) and Greenland (two and four species, respectively) (Danks 1981; Pollet 2015).
The empidoids occur on almost all main Archipelagos of the Arctic Ocean, with the only exception being
Franz Josef Land. We did not find any material from Severnaya Zemlya, however, Chernov (2004) noted some
unidentified Empididae from Bolshevik Island of this Archipelago. In addition, the empidoids were found from six
separate islands (from west to east)—Kildin, Kolguev, Vaygach, Dikson, Wrangel, Kolyuchin.
Only one species of Empididae (R. longestylata) was recorded from Svalbard Archipelago and it occurs only on
Spitsbergen. Ten species of Empididae belonging to two genera (Clinocera—2, Rhamphomyia—8) and one species
of Brachystomatidae (Heleodromia) are known now from Novaya Zemlya. The empidoids appear to be represented
by only two species (R. erinacioides and R. ursinella) on the New Siberian Archipelago.
One or two species of Empididae (only Rhamphomyia) are known from separate small Arctic islands, including
Kildin, Kolguev, Vaygach, Dikson and Kolyuchin.
The empidoid fauna of Wrangel Island is surprisingly rich. It includes all three families of Empidoidea (except
Dolichopodidae) found in our study and Hybotidae are known only from this island: Brachystomatidae (1 genus,
2 species), Empididae (4 genera, 22 species), Hybotidae (2 genera, 3 species). These numbers may be somewhat
influenced by higher intensity of collecting efforts conducted on Wrangel Island compared to most other island
territories discussed in this article (probably only with the exception of Svalbard). Nevertheless, we believe that they
are close to the real pattern among the Eurasian Arctic islands. It should also be noted that the species richness of
the empidoid fauna of Wrangel Island is even significantly higher than that of the much more well-studied islands
of the Canadian-Greenland sector (Sinclair 1994; 2008; Sinclair et al. 2011, 2019).
Six species of Empididae and one species of Hybotidae are known currently only from the Russian Arctic
islands (Novaya Zemlya—2 species of Empididae; Wrangel—4 species of Empididae and 1 species of Hybotidae).
Among the remaining 30 species of empidoids, 22 (60% of all species discussed in this paper) are Holarctic in
distribution. Our study suggests that empidoids are probably among the most speciose groups of Diptera Brachycera
on the Arctic islands (Chernov 1995). More detailed discussions on the zoogeographical and ecological features of
empidoids of the Arctic islands will be presented in a forthcoming paper by the authors.
To conclude, we hope that this paper will assist in the understanding of our present-day knowledge of empidoid
flies from the Eurasian Arctic islands and form a starting point for further studies into the ecology and conservation
of these flies in this region.
Acknowledgements
We are indebted to Yngve Brodin (NHRS), Pekka Vilkamaa, Jere Kahanpää (MZH) and Geir Søli (NHMO), as
well as their respective institutions, for the possibility to work with the Empididae collections and for the loan of
specimens used in this study. Fedor Konstantinov (Department of Entomology, St. Petersburg State University,
Russia) kindly provided equipment and software for taking digital images. Alexey Kovalev (ZIN) helped with some
photos. The study of Igor Shamshev was performed in the frames of the Russian State Research Project no. АААА-
А19-119020690101-6 and was supported by the Russian Foundation for Basic Research (grant no. 18-04-00354А)
and the Presidium RAS program no.41 “Biodiversity of natural systems and biological sources of Russia”. The work
of Olga Khruleva was supported by the Presidium of the Russian Academy of Sciences programs “Biodiversity
and Natural Resources of Russia: Structural and Functional Organization of Ecosystems and Communities” and
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72 · Zootaxa 4848 (1) © 2020 Magnolia Press
“Evolution of the Organic World: the Role and Influence of Planetary Processes”. The authors are sincerely grateful
to A.R. Gruzdev, director of the “Wrangel Island” Nature Reserve, for his help in field investigations, and to I.P.
Oleinikov, L.F. Volkova (Simon), U.V. Babiy and A.A. Rodionov, staff of the Nature Reserve, for help in insect
collecting on Wrangel Island. Toyohei Saigusa (Fukuoka, Japan) sorted and illustrated the CNC Rhamphomyia
collection, which greatly assisted the identification of several species in this study. Miroslav Barták (Prague, Czech
Republic) and an anonymous reviewer kindly reviewed the manuscript and made important remarks.
References
Baer, K.E. (1838) Expédition à Novaia-Semlia et en Laponie. Thierisches Leben. Bulletin Scientfique publé par l’Académie
Impériale des Sciences de St.-Pétersbourg, III, 343–351.
Barták, M. (1981) A revision of the Rhamphomyia albosegmentata - group (Diptera, Empididae), with descriptions of new
species. Acta Universitatis Carolinae Biologica, 11, 361–407.
Barták, M. (1982) The Czechoslovak species of Rhamphomyia (Diptera, Empididae), with description of a new species from
central Europe. Acta Universitatis Carolinae Biologica, 1980 (1982), 381–461.
Barták, M. (1985) A revision of the Rhamphomyia subgenus Lundstroemiella (Diptera, Empididae), with description of a new
species. Acta Universitatis Carolinae Biologica, 1982–1984 (1985), 9–46.
Barták, M. (1999) Two new Europaean species of Rhamphomyia subgenus Lundstroemiella (Diptera: Empididae). Klapalekiana,
35, 95–102.
Barták, M. (2006) Three new West Palaearctic species of Rhamphomyia subgenus Lundstroemiella (Diptera, Empididae).
Biologia, 61, 503–508.
https://doi.org/10.2478/s11756-006-0083-z
Barták, M. (2007) Catalogue of the Palaearctic species of the Rhamphomyia subgenus Lundstroemiella (Diptera: Empididae),
with new distribution records. Acta Zoologica Universitatis Comenianae, 47 (2), 101–104.
Barták, M. (2015) 17.13 Empididae (Dance or Dagger flies). In: Böcher, J., Kristensen, N.P., Pape, T. & Vilhelmsen, L.
(Eds.), The Greenland Entomofauna. An identification manual of insects, spiders and their allies. Fauna Entomologica
Scandinavica, 44, pp. 575–576.
Barták, M. & Danielson, R. (2007) Revision of Rhamphomyia species (Diptera: Empididae) described by J.W. Zetterstedt. Acta
Zoologica Universitatis Comenianae, 47 (2), 105–114.
https://doi.org/10.5962/bhl.part.80397
Barták, M. & Kubík, S. (2009) Two new east Palaearctic Rhamphomyia (Pararhamphomyia) (Diptera: Empididae). Entomological
News, 120, 76–86.
https://doi.org/10.3157/021.120.0114
Becker, Th. (1900) Beiträge zur Dipteren–Fauna Sibiriens. Nordwest-Sibirische Dipteren. Acta Societatis Scientiarum Fennicae,
26 (9), 1–66, 2 pl.
Becker, Th. (1908) Zur Kenntnis der Dipteren von Zentral-Asien. I. Cyclorrhapha schizophora holometopa und Orthorrhapha
brachycera. Annuaire du Musée zoologique de L’Académie Impériale des Sciences de St.-Pétersbourg, 12 (1907), 253–
317.
Becker, Th. (1915) Orthorrhapha Brachycera. In: Becker, Th., Dziedzicki, H., Schnabl, J. & Villeneuve, J. (Eds.), Résultats
scientifiques de l’expédition des frères Kuznecov (Kouznetzov) a l’Oural arctique en 1909, sous la direction de H. Backlund.
Livr. 7. Diptera. Mémoires de L’Académie Impériale des Sciences, VIII Série, Classe Physico-Mathématique, 28 (7), pp.
53–60.
Bezzi, M. (1903) Orthorrhapha Brachycera. In: Becker, T., Bezzi, M., Bischof, J., Kertész, K. & Stein, P. (Eds.), Katalog der
Paläartischen Dipteren. Band II. Wesseleìnyi, Budapest, 396 pp.
https://doi.org/10.5962/bhl.title.5139
Boheman, C.H. (1866) Spetsbergens Insekt-fauna. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar, 22 (1865),
563–577.
Chernov, Yu. I. (1995) The order Diptera (Insecta) in the Arctic fauna. Zoologicheskiy Zhurnal, 74, 68–83. [In Russian with
English summary]
Chernov, Yu. I. (2004) Terrestrial animals of polar desert on plateau of the Devon Island (Canadian Arctic Archipelago).
Zoologicheskiy Zhurnal, 83, 604–614. [in Russian with English summary]
Chvála, M. (1975) The Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica,
3, 1–336.
Chvála, M. (1989) Monograph of northern and central European species of Platypalpus Macq. (Diptera, Hybotidae), with data
on the occurrence in Czechoslovakia. Acta Universitatis Carolinae Biologica, 32 (1988), 209–376.
Chvála, M. & Wagner, R. (1989) Empididae. In: Soós, A. & Papp, L. (Eds.), Catalogue of Palaearctic Diptera. Vol. 6.
Therevidae—Empididae. Elsevier Science Publishing, Amsterdam, pp. 228–336.
Cockerell, T.D.A. (1916) Two Diptera of the genus Rhamphomyia from Colorado. The Canadian Entomologist, 48, 123–124.
https://doi.org/10.4039/Ent48123-4
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 73
Collin, J.E., (1923) Diptera (Orthorrhapha Brachycera and Cyclorrhapha) from Spitsbergen and Bear Island. Results of the
Oxford University Expedition to Spitsbergen, 1921.—No. 22. Annals and Magazine of Natural History, Series 9, 11 (61),
116–123.
https://doi.org/10.1080/00222932308632830
Collin, J.E. (1961) Empididae. In: British flies. Vol. 6. University Press, Cambridge, 782 pp.
Coquillett, D.W. (1900) Papers from the Harriman Alaska Expedition. IX. Entomological results (3): Diptera. Proceedings of
the Washington Academy of Sciences, 2, 389–464.
Coulson, S.J. (2007) Terrestrial and freshwater invertebrate fauna of High Arctic Archipelago of Svalbard. Zootaxa, 1448 (1),
41–58.
https://doi.org/10.11646/zootaxa.1448.1.2
Coulson, S.J. & Refseth, D. (2004) The terrestrial and freshwater invertebrate fauna of Svalbard. In: Prestrud, P., Strøm, H. &
Goldman, H.V. (Eds.), A catalog of the terrestrial and marine animals of Svalbard. Norwegian Polar Institute, Tromsø, pp.
57–122.
Coulson, S.J., Convey, P., Aakra, K., Aarvik, L., Ávila-Jiménez, M.L., Babenko, A., Biersma, E.M., Boström, S., Brittain, J.E.,
Carlsson, A.M., Christoffersen, K., De Smet, W.H., Ekremj, T., Fjellberg, A., Füreder, L., Gustafsson, D., Gwiazdowicz,
D.J., Hansen, L.O., Holmstrup, M., Hullé, M., Kaczmarek, Ů., Kolicka, M., Kuklin, V., Lakka, H.-K., Lebedeva, N.,
Makarova, O., Maraldo, K., Melekhina, E., Ødegaard, F., Pilskog, H.E., Simon, J.C., Sohlenius, B., Solhøy, T., Søli, G.,
Stur, E., Tanasevitch, A., Taskaeva, A., Velle, G., Zawierucha, K. & Zmudczyńska-Skarbek, K. (2014) The terrestrial
and freshwater invertebrate biodiversity of the archipelagoes of the Barents Sea; Svalbard, Franz Josef Land and Novaya
Zemlya. Soil Biology & Biochemistry, 68, 440–470.
https://doi.org/10.1016/j.soilbio.2013.10.006
Cumming, J.M. & Wood, D.M. (2017) [Chapter] 3. Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J.
(Eds.), Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. South
African National Biodiversity Institute, Pretoria, pp. 89–133.
Danks, H.V. (1981) Arctic arthropods. A review of systematics and ecology with particular reference to the North American
fauna. Entomological Society of Canada, Ottawa, 608 pp.
Downes, J.A. (1970) The feeding and mating behaviour of the specialized Empidinae (Diptera): observations on four species of
Rhamphomyia in the high arctic and a general discussion. The Canadian Entomologist, 102 (7), 769–791.
https://doi.org/10.4039/Ent102769-7
Fabricius, O. (1780) Fauna Groenlandica. Impensis Ioannis Gottlob Rothe, Hafniae, 452 pp, 1 pl.
Frey, R. (1915) Diptera-Brachycera aus den arktischen Küstengegenden Sibiriens. Résultats scientifiques de l’expédition
polaire Russe en 1900–1903, sous la direction du Baron E. Toll, Section E: Zoologie. Vol. II. Livr. 10. Zapiski Rossiyskoy
Akademii Nauk po Fiziko-Matematicheskomu Otdeleniyu (Transactions of the Russian Academy of Sciences for Physical
and Mathematical Branch), Series 8, 29 (10), 1–35.
Frey, R. (1922) Vorarbeiten zu einer Monographie der Gattung Rhamphomyia Meig. (Dipt., Empididae). Notulae entomologicae,
2, 1–10 + 33–45 + 65–77.
Frey, R. (1935) Entomologische Ergebnisse der schwedischen Kamtschatka-Expedition 1920–1922. 36. Diptera Brachycera. 5.
Empididae, Micropezidae, Lauxaniidae, Chloropidae. Arkiv för Zoologi, 28A (10), 1–8.
Frey, R. (1950) Neue paläarktische Rhamphomyia-Arten nebst Bestimmungstabelle der Rhamphomyia-Subgenera. Notulae
entomologicae, 29 (1949), 91–119.
Frey, R. (1955a) 28. Empididae. In: Lindner, E. (Ed.), Die Fliegen der palaearktischen Region, 4 (4), 433–480, taf. 37–42 (Lfg.
181).
Frey, R. (1955b) 28. Empididae. In: Lindner, E. (Ed.), Die Fliegen der palaearktischen Region, 4 (4), 481–528, taf. 43–48 (Lfg.
183).
Frey, R. (1956) 28. Empididae. Die Fliegen der palaearktischen Region, 4 (4): 577–639, Taf. 55–57 (Lfg. 188).
Gorodkov, K.B. & Kovalev, V.G. (1969) 44. Fam. Empididae. In: Bey-Bienko, G.Y. (Ed.), Keys to Insects of the European Part
of the USSR. 5 (1). Nauka, Moscow/Leningrad, pp. 573–670. [in Russian, English translation: 1988, pp. 886–1025]
Grichanov, I.Ya. (2006) A checklist and keys to North European genera and species of Dolichopodidae (Diptera). In: Plant
Protection News Supplements.VIZR, St. Petersburg, pp. 1–120.
Grichanov, I.Ya. (2017) Alphabetic list of generic and specific names of predatory flies of the epifamily Dolichopodoidae
(Diptera). 2nd Edition. In: Plant Protection News. Supplements. Issue 23. VIZR, St. Petersburg, pp. 1–563.
https://doi.org/10.5281/zenodo.884863
Grichanov, I.Ya. & Khruleva, O.A. (2018) Fauna and ecology of Dolichopodidae (Diptera) from Wrangel Island Nature Reserve
(Chukotka, Russia). Nature Conservation Research, 3, 37–45.
https://doi.org/10.24189/ncr.2018.023
Grootaert, P., Shamshev, I. & Van de Velde, I. (2012) Flowers as hunting ground for Platypalpus vegrandis Frey, 1943 (Diptera,
Hybotidae, Tachydromiinae). Bulletin de la Société royale belge d’entomologie/Bulletin van de Koninklijke Belgische
Vereniging voor Entomologie, 147 (2011), 239–240.
Hackman, W. (1968) Notes on the arthropod fauna of Spitsbergen, II: 11. Diptera: Empididae, Acalyptratae families and
Scatophagidae. Annales Entomologici Fennici, 34, 93–98.
Haliday, A.H. (1833) Catalogue of Diptera occurring about Holywood in Downshire. Entomological Magazine, London, 1,
SHAMSHEV ET AL.
74 · Zootaxa 4848 (1) © 2020 Magnolia Press
147–180.
Henricksen, K.L. & Lundbeck, W. (1917) Grønlands Landarthropoder (Insecta et Arachnida Groenlandicae). Meddelelser om
Grønland, 22, 481–823. [issue published 1917, entire volume bound and dated 1918]
https://doi.org/10.5962/bhl.title.3932
Holmgren, A.E. (1869) Bidrag till kännedomen om Beeren Eilands och Spetsbergens insektfauna. Kongliga Svenska Vetenskaps-
Akademiens Handlingar, 8 (5), 1–55.
Holmgren, A.E. (1880) Novas species insectorum cura et labore A. E. Nordenskiöldii e Novaia Semlia coactorum. Beckmanniana,
Holmiae (Stockholm), 24 pp.
Holmgren, A.E. & Aurivillius, C. (1883). Insecta a viris doctissimis Nordenskiöld illum ducem sequentibus in insulis Waigatsch
et Novaja Semlia anno 1875 collecta. Entomologisk Tidskrift, 4, 139–194.
Jonassen, T., Andersen, T. & Kvifte, G.M. (2013) Empidoidea (Diptera) from Finnmark, northern Norway. Norwegian Journal
of Entomology, 60, 201–245.
Kahanpää, J. (2014) Checklist of the Empidoidea of Finland (Insecta, Diptera). In: Kahanpää, J. & Salmela, J. (Eds.), Checklist
of the Diptera of Finland. ZooKeys, 441, pp. 183–207.
https://doi.org/10.3897/zookeys.441.7154
Kevan, P.G. (1972) Insect pollination of High Arctic Flowers. Journal of Ecology, 60, 831–847.
https://doi.org/10.2307/2258569
Kholod, S.S. (2013) Zonation in the plant cover on the Wrangel Island: syntaxonomical approach. Vegetation of Russia, 23,
89–121. [in Russian]
https://doi.org/10.31111/vegrus/2013.23.89
Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum caracteribus,
differentiis, synonymis, locis. Tomus 1. Editio decima. Reformata. L. Salvii, Holmiae, 824 pp.
https://doi.org/10.5962/bhl.title.542
Lundström, C. & Frey, R. (1913) Beitrag zur Kenntnis der Dipterenfauna des nördl. europäischen Russlands. Acta Societatis pro
Fauna et Flora Fennica, 37 (10), 1–20.
Malloch, J.R. (1919) The Diptera collected by the Canadian Arctic Expedition, 1913–1918. (excluding the Tipulidae and
Culicidae). In: Anderson, R.M. (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, pp. 34–90.
Melander, A.L. (1902) A monograph of the American Empididae. Part 1. Transactions of the American Entomological Society,
28, 195–368.
Melander, A.L. (1928) Diptera, Fam. Empididae. In: Wytsman, P. (Ed.), Genera Insectorum, Fasc. 185, “1927”. Louis Desmet-
Verteneuil, Bruxelles, pp. 1–434.
Oldenberg, L. (1915) Über einige Rhamphomyia-Arten. Archiv für Naturgeschichte, 80A (9), 69–91. [1914]
Oldenberg, L. (1922) Sechs alpine Rhamphomyia-Arten. Deutsche Entomologische Zeitschrift, 1922, 339–347.
https://doi.org/10.1002/mmnd.48019220314
Pape, T., Blagoderov, V. & Mostovski, M.B. (2011) Order DIPTERA Linnaeus, 1758. In: Zhang Z.-Q. (Ed.), Animal biodiversity:
An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148 (1), pp. 222–229.
https://doi.org/10.11646/zootaxa.3148.1.42
Plant, A. (2005) Heleodromia irwini Wagner (Diptera: Empididae), an English boreo-alpine relict? British Journal of Entomology
and Natural History, 18, 263.
Plant, A.R., Jonassen, T., Grootaert, P., Meyer, H., Pollet, M. & Drake, M. (2017) The arrow points north—endemic areas and
post-Devensian assembly of the British Empidoidea fauna (Insecta: Diptera). Biological Journal of the Linnean Society,
120, 852–868.
https://doi.org/10.1093/biolinnean/blw011
Pollet, M. (2015) Dolichopodidae. In: Böcher, J., Kristensen, N.P., Pape, T. & Vilhelmsen, L. (Eds.), The Greenland Entomofauna.
An identification manual of insects, spiders and their allies. Fauna Entomologica Scandinavica, 44, 577–581.
Poppius, B., Lundström, C. & Frey, R. (1917) Dipteren aus dem Sarekgebiet. Naturwissenschaftliche Untersuchungen des
Sarekgebirges in Schwedisch-Lappland. 4. Zoologie, Lfg. 6, 665–696, taf. 10.
Ringdahl, O. (1933) Översikt avde i Sverige funna arterna av släktet Atalanta Meig. (Clinocera Meig.) inom fam. Empididae.
Entomologisk Tidskrift, 54, 277–282.
Sack, P. (1923) Report of the scientific results of the Norwegian expedition to Novaya Zemlya, 1921. 15. Diptera. A.W. Broggers
Boktrykkery, Christiania (Oslo), 10 pp.
Saigusa, T. (1964) Some new species of the genus Rhamphomyia from Japan, II (Diptera, Empididae). Sieboldia, 3, 221–256.
Saigusa, T. (2012) A new Asio-Nearctic subgenus of Rhamphomyia (Diptera: Empididae: Empidinae). The Canadian
Entomologist, 144, 291–322.
https://doi.org/10.4039/tce.2012.28
Shamshev, I.V. (2001a) Thirteen new species of the genus Empis Linnaeus (Diptera: Empididae) from Asiatic part of the
Palaearctic region. An International Journal of Dipterological Research, 12, 195–227.
Shamshev, I.V. (2001b) 57. Fam. Atelestidae, 55. Fam. Hybotidae, 53. Fam. Empididae. In: Key to the insects of Russian Far
East. Vol. VI. Diptera and Siphonaptera. Pt 2. Dal’nauka, Vladivostok, pp. 150–151 + 258–286 + 296–346.
Shamshev, I.V. (2016) An annotated checklist of empidoid flies (Diptera: Empidoidea, except Dolichopodidae) of Russia.
EMPIDOID FLIES OF RUSSIAN ARCTIC ISLANDS Zootaxa 4848 (1) © 2020 Magnolia Press · 75
Proceedings of the Russian Entomological Society, 87, 3–183.
Shamshev, I.V. & Barkalov, A.V. (2009) Fauna and zoogeographic characteristic of dance flies (Diptera, Empididae) of Mountain
Altay. Euroasian Entomological Journal, 8, 319–323.
Shamshev, I.V. & Sinclair, B.J. (2018) Revision of Rhamphomyia species (Diptera: Empididae) described by A.E. Holmgren
from Novaya Zemlya Islands. Russian Entomological Journal, 27 (3), 307–314.
https://doi.org/10.15298/rusentj.27.3.13
Sinclair, B.J. (1994) Revision of the Nearctic species of Trichoclinocera Collin (Diptera: Empididae; Clinocerinae). The
Canadian Entomologist, 126, 1007–1059.
https://doi.org/10.4039/Ent1261007-4
Sinclair, B.J. (1999) Review of the Holarctic Clinocera appendiculata complex (Dipt., Empididae, Clinocerinae). Entomologist’s
Monthly Magazine, 135, 223–232.
Sinclair, B.J. (2008) The Systematics of New World Clinocera Meigen (Diptera: Empididae: Clinocerinae). NRC Research
Press, Ottawa, viii+245 pp.
Sinclair, B.J. (2017) [Chapter] 53. Brachystomatidae. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.), Manual of Afrotropical
Diptera. Vol. 2. Nematocerous Diptera and lower Brachycera. Suricata 5. South African National Biodiversity Institute,
Pretoria, pp. 1251–1256.
Sinclair, B.J., Brooks, S.E., Cumming, J.M. & Coovert, G.A. (2011) Revision of the Nearctic species of Heleodromia (Diptera:
Empidoidea: Brachystomatidae). The Canadian Entomologist, 143, 629–651.
https://doi.org/10.4039/n11-036
Sinclair, J.B., Vajda, É.A., Saigusa, T., Shamshev, I.V. & Wheeler, T.A. (2019) Rhamphomyia Meigen of the Canadian Arctic
Archipelago, Greenland and Iceland (Diptera: Empididae). Zootaxa, 4670 (1), 1–94.
https://doi.org/10.11646/zootaxa.4670.1.1
Yang, D., Zhang, K., Yao, G. & Zhang, J. (2007) World Catalog of Empididae (Insecta: Diptera). China Agricultural University
Press, Beijing, 599 pp.
Wagner, R. (1985) A revision of the genus Heleodromia (Diptera, Empididae) in Europe. Aquatic Insects, 7, 33–43.
https://doi.org/10.1080/01650428509361196
Wahlberg, E. & Johanson, K.A. (2018) Molecular phylogenetics reveals novel relationships within Empidoidea (Diptera).
Systematic Entomology, 43, 610–636.
https://doi.org/10.1111/syen.12297
Zetterstedt, J.W. (1838) Sectio tertia. Diptera. Dipterologia Scandinaviae amicis et popularibus carissimus. In: Insecta Lapponica.
L. Voss, Lipsiae (Leipzig), pp. 477–868.
Zetterstedt, J.W. (1842) Diptera Scandinaviae disposita et descripta. Tomus primus. Officina Lundbergiana, Lundae (Lund),
xvi + 440 pp.
Zetterstedt, J.W. (1859) s.n. In: Diptera Scandinaviae disposita et descripta. (Part.) Tomus tridecimus [sic!] seu Supplementum
Quartum, continens addenda, corrigenda & emendanda tomis duodecim prioribus. una cum conspectus omnium generum.
ex Officina Lundbergiana, sumtibus auctoris, Lundae (Lund), pp. I–XVI + 4943–6190.
