Research Article |
Corresponding author: Jindřich Roháček ( rohacek@szm.cz ) Academic editor: Marc De Meyer
© 2022 Jindřich Roháček, Andrey A. Przhiboro.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Roháček J, Przhiboro AA (2022) Pullimosina (Pullimosina) turfosa sp. nov. and other Sphaeroceridae (Diptera) from peat bogs in the North Caucasus (Russia). ZooKeys 1132: 1-49. https://doi.org/10.3897/zookeys.1132.94579
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The first data about Sphaeroceridae occurring on eight montane and foothill peat bogs of various types in the North Caucasus (Russia) are presented. A total of 38 species has been recorded and their affinity to peat-bog habitats is discussed. A single species is classified as a tyrphobiont, viz. the strongly brachypterous Pullimosina (Pullimosina) turfosa sp. nov. being strictly associated with Sphagnum hummocks in peat bogs. This new species is described and illustrated in detail and its relationships, biology, and wing reduction are discussed. Only three species are considered tyrphophilous or probably tyrphophilous, viz. Ischiolepta nitida (Duda, 1920), Phthitia (Kimosina) longisetosa (Dahl, 1909), and Spelobia ibrida Roháček, 1983. The majority of recorded species do not have close affinity to peat bogs and are treated as tyrphoneutral, and Rachispoda hostica (Villeneuve, 1917) is probably tyrphoxenous due to occasional occurrence in a peat bog. Species composition of Sphaeroceridae on Caucasian peat bogs is discussed in comparison to those known from peat bogs in other parts of Europe. Taxonomic notes are given on Minilimosina (Svarciella) species of the M. vitripennis group. Six species (including P. turfosa sp. nov.) are new additions to the fauna of Russia.
biology, brachyptery, distribution, lesser dung flies, Limosininae, relationships, taxonomy, tyrphophily
Sphaeroceridae or Lesser dung flies are a relatively large family of Acalyptratae, with more than 1,800 described (and at least 5,000 estimated) species in the world fauna of Diptera (
Insect fauna of mires (peat bogs in a broad sense, i.e., peatlands where peat is currently being formed and accumulating) has been relatively well studied both in Europe and North America (for reviews, see
This study is devoted to Sphaeroceridae obtained from montane and submontane peat bogs in the North Caucasus by the second author. These mires are rare, mostly small and isolated habitats situated near the southernmost limit of the occurrence of this habitat type in Europe and the whole of the Palaearctic Region. They are characterized by specific environmental conditions and distinctive composition of vegetation (
A total of 119 adults of Sphaeroceridae was collected in peat bogs together with other macroinvertebrates in the scope of faunal and ecological studies of these mires in early May 2016, late May to early June 2018, and in September 2018. Most of the examined material is deposited in Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia (
On each mire, most sampling effort was focused on collecting macroinvertebrates from dry and wet habitats where Sphagnum mosses dominate or are abundant. The following main sampling techniques were used in each mire: sweep-netting with aerial net over grassy vegetation (in different daytime periods and weather), yellow pan traps, pitfall traps, sifting substrate of Sphagnum cushions or hummocks located in drier places, trampling Sphagnum cushions located in wet places (like shorelines of in-mire lakelets), sweep-netting with aquatic net in puddles and along shorelines of lakelets, and sampling the substrate (Sphagnum, other plants, turf, and litter). The latter samples were washed in sieves (the smallest 0.25 mm mesh), then macroinvertebrates were extracted by flotation in a strong solution of NaCl combined with hand-sorting of the coarse fraction. However, most Sphaeroceridae were collected by sweep-netting; only a few specimens were obtained by sifting, yellow pan traps, and from the samples of substrata (indicated below as “sample no.”).
Abdomens of some specimens were detached, cleared by boiling for several minutes in 10% solution of potassium hydroxide (KOH) in water, then neutralized in 10% solution of acetic acid (CH3COOH) in water, washed in water and subsequently transferred to glycerin. Postabdominal structures were dissected and examined in a drop of glycerin under binocular microscopes (Reichert, Olympus). Detailed examinations of genital structures were performed with a compound microscope (Zeiss Jenaval). After examination, all dissected parts were put into small plastic tubes containing glycerin, sealed with hot forceps and pinned below the respective specimens. Specimens with abdomen removed and terminalia dissected are indicated in the list of material by the abbreviation “genit. prep.”
Legs and details of the male and female genitalia were drawn by means of an Abbe’s drawing apparatus on a compound microscope (Zeiss Jenaval) at magnification 130–500×. Wings were photographed on an Olympus BX51 compound microscope with an attached digital camera (Canon EOS 1200D). Whole adult (dry-mounted) specimens and heads were photographed by means of a Canon EOS 5D Mark III digital camera with a Nikon CFI Plan 10×/0.25NA 10.5mm WD objective attached to a Canon EF 70–200mm f/4L USM zoom lens. The specimen photographed by means of the latter equipment was repositioned upwards between each exposure using a Cognisys StackShot Macro Rail and the final photograph was compiled from multiple layers (~ 40) using Helicon Focus Pro 7.0.2. The final images were edited in Adobe Photoshop CS6. All morphological illustrations were prepared by the first author.
Five characteristics of the new species were measured: body length (measured from anterior margin of head to end of cercus, thus excluding the antenna), index t2: mt2 (= ratio of length of mid tibia: length of mid basitarsus), wing length (from wing base to wing tip), wing width (maximum width), Cs1: Cs2 (= ratio of length of 1st costal sector: length of 2nd costal sector). All type specimens were measured.
Label data of the type specimens are presented strictly verbatim including information on form and color of all associated labels. Locality data of other specimens examined are given in brief form because all other information is given under the descriptions of localities. Localities are listed in the same order as in their descriptions below. Biological information obtained from the material examined and literature are given in the Comments paragraph. General distribution of species recorded are based on
The affinity of a species to peat-bog habitat has been judged by the first author based on his knowledge of the biology, autecology and distribution of the species. Four categories are differentiated according to the degree of association with bog habitats following
These categories can be compared with those more generally used in North Europe (e.g., by
Morphological terminology follows that used for Sphaeroceridae by
A1 anal vein
ac acrostichal (seta)
ads additional (setulae) on frons
asc additional sclerite
C costa
ce cercus
Cs1, Cs2 1st, 2nd costal sector
CuA1 cubitus
cx1 fore coxa
dc dorsocentral (seta)
dp distiphallus
ea ejacapodeme
ep epandrium
f1, f2, f3 fore, mid, hind femur
g genal (seta)
gs gonostylus
h humeral cross-vein
hu humeral (= post-pronotal) (seta)
hy hypandrium
ifr interfrontal (seta)
M media
ma medandrium
mt2, mt3 mid, hind basitarsus
oc ocellar (seta)
occe outer occipital (seta)
occi inner occipital (seta)
ors fronto-orbital (seta)
pg postgonite
pha phallapodeme
pp phallophore
prg pregonite
pvt postvertical (seta)
R1 1st branch of radius
R2+3 2nd branch of radius
R4+5 3rd branch of radius
S1–S10 abdominal sterna
sc scutellar (seta)
stpl sternopleural (= kat-episternal) (seta)
T1–T10 abdominal terga
t1, t2, t3 fore, mid, hind tibia
va ventroapical seta on t2
vi vibrissa
vte outer vertical (seta)
vti inner vertical (seta)
For mire types and characteristics, we mostly follow the terminology and definitions adopted by
Eight montane and submontane mires (peat bogs) were studied, all situated in the central part of the northern slope of the Greater Caucasus Range (Fig.
The mires under study strongly differ in the size (0.0004 to ca. 0.5 km2), altitude (800 to 2290 m), trophic status (oligo- to eutrophic), origin, and type (Sphagnum-, Carex-Sphagnum- and Carex-dominated). Conditions of the study mires and previous publications about these mires were briefly reviewed in
Three small transitional mires called here Kurnoyatsu-1, Kurnoyatsu-2, and Kurnoyatsu-3 (having no official names) are situated on the slope at left bank of the Kurnoyatsu River, ca. 3 km SE of Verkhnyaya Balkaria Village. These mires are situated in the alpine zone near the forest edge, on small flat terraces at different heights, 200–500 m from each other. The mires are limnogenous; each has a lakelet in the middle, with swing moor at shoreline. Relief of hummocks is not developed. Kurnoyatsu-1 (Fig.
Region (area within) | Bog name | Coordinates | Altitude (m) | Mire area (m2)* | Trophic status | Habitats | Dominating species of Sphagnum | Dominating species of phanerogams |
---|---|---|---|---|---|---|---|---|
Kabardino-Balkaria (eastern Balkaria) | Kurnoyatsu-1 | 43.10062°N, 43.48418°E | 1776 | 2600 | oligotrophic to meso-oligotrophic | dry | S. magellanicum-coll. (S. divinum) | Molinia caerulea, Menyanthes trifoliata |
wet (near lake) | S. squarrosum, S. teres, S. flexuosum, S. obtusum | Carex rostrata, C. canescens, C. ? diandra, Menyanthes trifoliata | ||||||
Kurnoyatsu-2 | 43.09834°N, 43.47776°E | 1810 | 4000 | oligotrophic to meso-oligotrophic | dry | S. fuscum | Rhododendron luteum, Empetrum nigrum, Carex rostrata | |
wet (near lake) | S. fallax, S. flexuosum | Carex rostrata, C. canescens, Calamagrostis sp. | ||||||
Kurnoyatsu-3 | 43.09714°N, 43.47950°E | 1836 | 1500 | oligotrophic to meso-oligotrophic | mostly wet | S. fallax, S. flexuosum | Carex rostrata, Eriophorum angustifolium | |
Ushtulu | 42.97457°N, 43.33263°E | 1995 | 180000 | eutrophic | dry | S. warnstorfii | Carex rostrata | |
wet | S. warnstorfii | Carex rostrata | ||||||
North Ossetia-Alania (western Digoria) | Chifandzar | 42.91867°N, 43.51493°E | 2289 | 520000 | eutrophic | dry (large hummocks) | S. teres | Carex rostrata, C. sp., Nardus stricta |
wet | S. subsecundum | Carex rostrata, Nardus stricta | ||||||
Kubus-larger | 42.89350°N, 43.57733°E | 2077 | 2300 | oligotrophic | dry | S. capillifolium | Molinia caerulea, Eriophorum angustifolium, Nardus stricta | |
Kubus-smaller | 42.89350°N, 43.57733°E | 2080 | 400 | oligotrophic | mostly wet | S. subsecundum | Carex rostrata, C. magellanica, C. lasiocarpa, Nardus stricta | |
North Ossetia-Alania (Tarskaya Hollow) | Tarskoe | 42.96311°N, 44.72636°E | 800 | 62000 | meso-oligotrophic | dry | S. magellanicum-coll. (S. divinum), S. centrale | Molinia caerulea |
wet (near ditches) | S. subsecundum | Molinia caerulea, Carex sp. |
Ushtulu mire (also called “Narzannoe”; Fig.
Chifandzar mire (Fig.
Two small transitional mires of Kubus, Kubus-larger (Fig.
Tarskoe peatland (Fig.
Three Kurnoyatsu mires are occasionally used for grazing horses, while Chifandzar and Tarskoe mires, for grazing cattle. Chifandzar and Kubus mires are located within the Alania National Park; Tarskoe peatland has a formal protection status as a regional natural monument; Ushtulu mire is situated within the borders of the Kabardino-Balkaria State High-Mountain Reserve; Kurnoyatsu mires have no protection status.
Holotype
♂ labelled: “Russia: N Ossetia, W Digoria, Chifandzar mire in Kharesidon River valley, 42.91867°N, 43.51493°E, 2289 m, sifting from Sphagnum teres hummocks, 18.ix.2018, A. Przhiboro leg.“, ”Holotypus ♂ Pullimosina (Pullimosina) turfosa sp. n., J. Roháček det. 2022“ (red label). The specimen is dried from ethanol and mounted on pinned triangular card, intact (deposited in
This strongly brachypterous species is named turfosa (= peaty, Latin adjective) owing to its strict association with Sphagnum hummocks in the type locality.
Male (Figs
Head
(Figs
Thorax
brown to pale brown (pleuron paler) and greyish brown microtomentose; mesonotum subshining, pleuron and scutellum more densely microtomentose and duller (Figs
Legs
brown to pale brown, coxae, trochanters and knees ochreous to yellow; fore coxa and all trochanters lightest, dirty yellow. Chaetotaxy: f1 with a posterodorsal row of 6 or 7 shorter setae and a posteroventral row of 7 or 8 longer setae in addition to ventrobasal fine seta (Fig.
Wing
(Figs
Abdomen
(Figs
Pullimosina (Pullimosina) turfosa sp. nov. (paratypes) 18 male fore leg without tarsus, posteriorly 19 male mid tibia, dorsally 20 male mid femur, tibia and basitarsus, anteriorly 21 female mid tibia and basitarsus, anteriorly 22 male right wing, dorsally. Scale bars: 0.2 mm (18–21); 0.1 mm (22).
Pullimosina (Pullimosina) turfosa sp. nov. (male paratype) 23 S5 and postabdominal sclerites, ventrally 24 gonostylus, laterally 25 external genitalia, caudally 26 ditto, laterally. Scale bars: 0.1 mm (23, 25, 26); 0.05 mm (24). Abbreviations: ce – cercus, dp – distiphallus, ep – epandrium, gs – gonostylus, hy – hypandrium, ma – medandrium, pg – postgonite, pha – phallapodeme, S – sternum.
Genitalia
: Epandrium (Figs
Female (Figs
Postabdomen
(Figs
Pullimosina (Pullimosina) turfosa sp. nov. (male paratype) 27 aedeagus (phallus) dorsally 28 ditto, laterally 29 aedeagal complex, laterally 30 pregonite (enlarged), laterally. Scale bars 0.05 mm (27-29), 0.02 mm (30). Abbreviations: dp – distiphallus, ea – ejacapodeme, pg – postgonite, pha – phallapodeme, pp – phallophore, prg – pregonite.
Despite a number of peculiarities in the male and female terminalia and unusual reduction of wing venation, Pullimosina turfosa sp. nov. clearly is a representative of the subgenus Pullimosina s. str. (
Pullimosina turfosa can be most easily recognized from all Holarctic Pullimosina species by its strongly abbreviated wings with very characteristic venation (Figs
Pullimosina (Pullimosina) turfosa sp. nov. (female paratype) 33 spermathecae 34 postabdomen, dorsally 35 ditto, ventrally 36 ditto, laterally 37 S8 and additional sclerite, ventrally 38 spectacles-shaped sclerite, laterally 39 ditto and single spermatheca, ventrally. Scale bars: 0.1 mm (34–36); 0.05 mm (33, 37–39). Abbreviations: asc – additional sclerite, ce – cercus, S – sternum, T – tergum.
The peculiar reduction of the wing and its veins in P. turfosa (Fig.
The species is known only from its type locality in Russia, North Ossetia (Caucasus Mts).
All specimens of the new species were collected on 17 and 18 August 2018 in a high-montane Chifandzar mire (Fig.
All type specimens but one were collected from large Sphagnum hummocks (Fig.
Most specimens were collected by means of sifting substrata of hummocks. Two females were collected by washing and subsequent flotation of substrate in NaCl solution: one specimen was sampled from the same habitat, and another one, from moist substrate beyond the hummocks, with predominating Sphagnum subsecundum and Carex rostrata. Hence, most individuals of P. turfosa concentrate in hummocks but some flies may also occur at some distance from them. No specimens were collected in early summer (2–3 June), suggesting that the adults of P. turfosa appear later.
Due to exclusive association of P. turfosa with the sphagnetum habitat, particularly with hummocks, we consider it a tyrphobiont (= eucoenic to peat-bog habitat) sphagnicolous species. Interestingly, no specimens of P. turfosa were collected from similar substrata in other bogs using the same techniques (sifting and washing/flotation). It is possible that the new peculiar species is confined to high montane bogs or even endemic to Chifandzar, considering that the montane bogs of the North Caucasus are rare and isolated island ecosystems.
Borborillus uncinatus (Duda, 1923)
Borborillus vitripennis (Meigen, 1830)
Copromyza equina Fallén, 1820
Lotophila atra (Meigen, 1830)
Norrbomia costalis (Zetterstedt, 1847)
Norrbomia sordida (Zetterstedt, 1847)
Ischiolepta nitida (Duda, 1920)
Chaetopodella scutellaris (Haliday, 1836)
Coproica acutangula (Zetterstedt, 1847)
Coproica ferruginata (Stenhammar, 1855)
Coproica lugubris (Haliday, 1836)
Eulimosina ochripes (Meigen, 1830)
Gonioneura spinipennis (Haliday, 1836)
Leptocera fontinalis (Fallén, 1826)
Leptocera nigra Olivier, 1813
Leptocera oldenbergi (Duda, 1918)
Minilimosina (Minilimosina) fungicola (Haliday, 1836)
Minilimosina (Minilimosina) gemella Roháček, 1983
Minilimosina (Minilimosina) sp.
Minilimosina (Svarciella) pujadei Carles-Tolrá, 2001
Minilimosina (Svarciella) vitripennis (Zetterstedt, 1847)
Opacifrons coxata (Stenhammar, 1855)
Opalimosina (Opalimosina) mirabilis (Collin, 1902)
Opalimosina (Pappiella) liliputana (Rondani, 1880)
Phthitia (Kimosina) longisetosa (Dahl, 1909)
Pseudocollinella (Spinotarsella) humida (Haliday, 1836)
Pullimosina (Pullimosina) heteroneura (Haliday, 1836)
Pullimosina (Pullimosina) turfosa sp. nov.
Rachispoda hostica (Villeneuve, 1917)
Rachispoda lutosoidea (Duda, 1938)
Spelobia clunipes (Meigen, 1830)
Spelobia czizeki (Duda, 1918)
Spelobia ibrida Roháček, 1983
Spelobia luteilabris (Rondani, 1880)
Spelobia parapusio (Dahl, 1909)
Spelobia rufilabris (Stenhammar, 1855)
Spelobia talparum (Richards, 1927)
Terrilimosina schmitzi (Duda, 1918)
Borborillus uncinatus (Duda, 1923) – TN
Material. Kubus-larger, sweep-netting, 9.v.2016, 1♂.
Comments. A largely coprophagous species, widespread in temperate and northern belt of the Palaearctic Region. Its occurrence on a peat bog is surely occasional due to attraction to some mammal excrement. There is a single previous record from two bogs in Wales (
Borborillus vitripennis (Meigen, 1830) – TN
Material. Kubus-larger, sweep-netting, 9.v.2016, 1♂.
Comments. A coprophagous Palaearctic species, mainly associated with horse dung on pastures. There are only scarce records from peat bogs in England (
Copromyza equina Fallén, 1820 – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 2♂.
Comments. A subcosmopolitan coprophagous species, occurring on (preferably horse) dung on pastures but also on manure. Also known to sometimes occur on peat bogs in Great Britain (
Lotophila atra (Meigen, 1830) – TN
Material. Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 1♀; wet habitat (lake margin), 7.vi.2018, 1♀. Tarskoe, sweep-netting, 10.v.2016, 1♀; same but 1.vi.2018, 1♂; same but 11.ix.2018, 2♂.
Comments. A Holarctic, predominantly coprophagous, species, common on various animal dung. Although repeatedly recorded from several peat bogs in Europe (e.g.,
Norrbomia costalis (Zetterstedt, 1847) – TN
Material. Chifandzar, sweep-netting 2 (daytime/sun), 18.vi.2018, 1 specimen (damaged, without abdomen); sweep-netting 1 (evening/rain), 17.ix.2018, 1♂ 1♀ (1♂ dry, genit. prep.).
Comments. A West Palaearctic coprophagous species, associated with (preferably horse) excrement on pastures. Formerly only recorded from a peat bog in Wales (
Norrbomia sordida (Zetterstedt, 1847) – TN
Material. Tarskoe, sweep-netting, 10.v.2016, 1♂ (dry, genit. prep.).
Comments. Originally a Holarctic (introduced to Neotropical, Oriental, and Oceanian Regions) coprophagous species, mainly occurring on dung of hoofed animals. There is no previous record from peat bogs.
Ischiolepta nitida (Duda, 1920) – TPH?
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♂ (genit. prep.).
Comments. A Palaearctic saprophagous (mainly coprophagous) species preferentially occurring in humid places in open and woodland habitats at higher altitudes. It seems to have distinct affinity to bog habitats and was recorded from peat bogs in England (
Chaetopodella scutellaris (Haliday, 1836) – TN
Material. Tarskoe, sweep-netting, 11.ix.2018, 1♂.
Comments. A common, largely coprophagous species, widespread in the Palaearctic Region, most common on dung on pastures. Although it is known to occur on various peat bogs (
Coproica acutangula (Zetterstedt, 1847) – TN
Material. Kurnoyatsu-1, dry habitat (Sphagnum magellanicum), sweep-netting, 6.vi.2018, 1♀; same but 22.ix.2018, 2♀.
Comments. A widespread (subcosmopolitan) coprophagous species associated with (mainly horse) dung on pastures. Records from bog habitats are very sporadic (
Coproica ferruginata (Stenhammar, 1855) – TN
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♂.
Comments. A cosmopolitan coprophagous species, very common on dung. Although repeatedly recorded from peat bogs and similar mire habitats in Europe (e.g.,
Coproica lugubris (Haliday, 1836) – TN
Material. Tarskoe, sweep-netting, 1.vi.2018, 1♀.
Comments. Another widespread (in the Palaearctic and Oriental Regions) coprophagous species, common on dung in pastures. It has been only occasionally recorded from peat bogs in Europe (
Eulimosina ochripes (Meigen, 1830) – TN
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♀. Tarskoe, sweep-netting, 11.ix.2018, 1♀.
Comments. A Holarctic phytosaprophagous species, mainly living on meadows. There are records from lagg meadows of peat bogs in England (
Gonioneura spinipennis (Haliday, 1836) – TN
Material. Kurnoyatsu-1, dry habitat (Sphagnum magellanicum), sweep-netting, 6.vi.2018, 1♀; same habitat, yellow pan traps, 6-8.vi.2018, 1♂; wet habitat (lake margin), sweep-netting, 6.vi.2018, 1♂; same but 7.vi.2018, 1♂. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♂. Chifandzar, wet habitat (Sphagnum subsecundum), sample Ч 2, 3.vi.2018, 1♀. Tarskoe, sweep-netting, 10.v.2016, 1♂ 1♀.
Comments. A common Holarctic polysaprophagous species, mainly living on dung. In Central Europe it is relatively frequent on peat bogs (
Leptocera fontinalis (Fallén, 1826) – TN
Material. Kurnoyatsu-1, dry habitat (Sphagnum magellanicum), sweep-netting, 6.vi.2018, 1♀; wet habitat (lake margin), sweep-netting, 6.vi.2018, 2♀. Kubus-smaller, sweep-netting (2), 14.ix.2018, 1♀.
Comments. A Holarctic saprophagous species, occurring in various humid habitat, mainly on decaying vegetation. Only
Leptocera nigra Olivier, 1813 – TN
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♂. Ushtulu, sweep-netting, 21.ix.2018, 1♀. Kubus-larger, sweep-netting, 4.vi.2018, 1♂; same but 14.ix.2018, 3♂ 2♀. Kubus-smaller, sweep-netting, 4.vi.2018, 1♂; same but 12.ix.2018, 2♂.
Comments. A very common species, widespread in the Old World but also introduced to Venezuela in the Neotropical Region (
*Leptocera oldenbergi (Duda, 1918) – TN
Material. Kurnoyatsu-1, dry habitat (Sphagnum magellanicum), sweep-netting, 6.vi.2018, 1♀ (genit. prep.). Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♀.
Comments. A rare species known from temperate Europe (Belgium, Czech Republic, Denmark, Germany, Great Britain, Hungary, Ireland, Latvia, Netherlands, Slovakia, Sweden, Switzerland) but also recorded from Georgia (
*Minilimosina (Minilimosina) fungicola (Haliday, 1836) – TN
Material. Kurnoyatsu-3, Sphagnum/sedge floating shores of lake, sweep-netting, 7.vi.2018, 2♀ (genit. prep.).
Comments. A Holarctic polysaprophagous species, habitat- and altitude-tolerant. It was recorded from mire habitats in England (
*Minilimosina (Minilimosina) gemella Roháček, 1983 – TN?
Material. Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 1♂ 1♀ (genit. prep.).
Comments. Also Holarctic, but essentially a Boreo-montane species, with a few records from Europe (cf.
Minilimosina (Minilimosina) sp.
Material. Chifandzar, sweep-netting 2, 18.vi.2018, 1♀ (genit. prep.).
Comments. Based on structures of the female postabdomen, this specimen cannot be associated with any of Minilimosina (s. str.) species known from Europe. It could either be conspecific with some of the poorly characterized species described by
*Minilimosina (Svarciella) pujadei Carles-Tolrá, 2001 – TN?
Material. Kubus-larger, sweep-netting, 9.v.2016, 1♂ (dry, genit. prep.).
Comments. Apart from the above new Pullimosina species, this species is the most surprising finding from the Caucasian peat bogs. It was described from a single male captured in a Malaise trap near a forest and a river at 1050 m in Andorra (
Taxonomic notes. Owing to this surprising record, the Caucasian specimen has been compared to a male of M. pujadei from the Czech Republic (redescribed by
Minilimosina (Svarciella) vitripennis (Zetterstedt, 1847) – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 1♀ (dry, genit. prep.). Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 1♂ (genit. prep.); same habitat, yellow pan traps, 7-8.vi.2018, 1♂ (genit. prep.). Kurnoyatsu-3, Sphagnum/sedge floating shores of lake, sweep-netting, 7.vi.2018, 1♂. Kubus-larger, sweep-netting, 4.vi.2018, 1♂ 2♀.
Comments. A Holarctic polysaprophagous species associated with various, mainly open, habitats (meadows, pastures etc.). There are also a few records from mire habitats in England (
Heads of Minilimosina (Svarciella) species, frontally 40 M. (S.) bartaki Roháček, 2010, male paratype (Czech Republic) 41 M. (S.) pujadei Carles-Tolrá, 2001, male (Caucasus) 42 M. (S.) vitripennis (Zetterstedt, 1847), female (Czech Republic) 43 Ditto, female (Caucasus). Scale bar 0.2 mm.
Taxonomic notes. A female with enlarged glabrous frontal triangle (Fig.
Opacifrons coxata (Stenhammar, 1855) – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 1♀; same habitat, yellow pan traps, 6-8.vi.2018, 1♂. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♀. Ushtulu, sweep-netting, 21.ix.2018, 1♂.
Comments. A common Palaearctic species associated with mud on shores of water bodies and in marshland habitats. It is eurytopic and, therefore, able to live on peat mud in mires (see
Opalimosina (Opalimosina) mirabilis (Collin, 1902) – TN
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♀.
Comments. A subcosmopolitan ubiquitous, predominantly coprophagous species. It is extremely habitat-tolerant and, therefore, can also occur on various excrement on peat bogs; see e.g.,
Opalimosina (Pappiella) liliputana (Rondani, 1880) – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 1♂ 1♀ (genit. prep.). Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 1♂.
Comments. A Holarctic and widely habitat-tolerant species developing in various decaying substrates. In Europe, it was sporadically recorded also from mire habitats, viz. by
Phthitia (Kimosina) longisetosa (Dahl, 1909) – TPH?
Material. Chifandzar, dry habitat (Sphagnum teres hummocks), sifting, 18.ix.2018, 1♀ (genit. prep.).
Comments. A rather rare West Palaearctic species with poorly known biology, usually occurring in damp meadows, fens and humid forests, sometimes in burrows of small mammals (cf.
Pseudocollinella (Spinotarsella) humida (Haliday, 1836) – TN
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 24.ix.2018, 1♀. Kubus-smaller, sweep-netting, 4.vi.2018, 1♀.
Comments. A common paludicolous species, widespread in the Palaearctic Region. It is associated with muddy habitats and is also known to live on peat bogs in Europe (
Pullimosina (Pullimosina) heteroneura (Haliday, 1836) – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 1♂. Tarskoe, 10.v.2016, sweep-netting, 1♀.
Comments. A cosmopolitan, eurytopic polysaprophagous species, common on various rotting substrates but records (of only single specimens) from peat bogs are scarce (
*Pullimosina (Pullimosina) turfosa sp. nov. – TB
Material. Chifandzar, dry habitat (Sphagnum teres hummocks), sample Ч 14, 17.ix.2018, 1♀; same habitat, sifting, 18.ix.2018, 6♂ 5♀; wet habitat (Sphagnum subsecundum), sample Ч 9, 17.ix.2018, 1♀ (see also type material above; some with genit. prep.)
Comments. This new terricolous and strongly brachypterous species (described above) is surely tyrphobiont, i.e., exclusively associated with peat-bog habitat, because living in Sphagnum hummocks (Fig.
Rachispoda hostica (Villeneuve, 1917) – TX?
Material. Kurnoyatsu-2, wet habitat (lake margin), sweep-netting, 7.vi.2018, 1♀ (genit. prep.).
Comments. An uncommon Palaearctic species (but unrecorded from most of Western Europe and North Africa) with easternmost records from Mongolia (
Rachispoda lutosoidea (Duda, 1938) – TN
Material. Tarskoe, sweep-netting, 30.v.2018, 1♂.
Comments. A common West Palaearctic species (also recorded from the Caucasus, cf.
Spelobia clunipes (Meigen, 1830) – TN
Material. Kurnoyatsu-1, wet habitat (lake margin), sweep-netting, 6.vi.2018, 2♀. Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 3♀; wet habitat (lake margin), sweep-netting, 7.vi.2018, 2♀; wet Sphagnum habitat (lake margin), yellow pan traps, 7-8.vi.2018, 2♂. Kurnoyatsu-3, Sphagnum/sedge floating shores of lake, sweep-netting, 7.vi.2018, 1♂ 2♀ (genit. prep.). Kubus-larger, sweep-netting, 4.vi.2018, 2♂; sweep-netting (2), 14.ix.2018, 1♀. Tarskoe, sweep-netting, 10.v.2016, 6♂ 5♀ (genit. prep.).
Comments. A very common, eurytopic and polysaprophagous Holarctic species. Because of its wide habitat and trophic tolerance, it was often recorded from peat bogs in Europe (e.g.,
Spelobia czizeki (Duda, 1918) – TN
Material. Kurnoyatsu-1, dry habitat (Sphagnum magellanicum), sweep-netting, 6.vi.2018, 1♂ (genit. prep.). Kurnoyatsu-3, Sphagnum/sedge floating shores of lake, sweep-netting, 7.vi.2018, 1♀.
Comments. A rather uncommon polysaprophagous Palaearctic species associated with various subterranean habitats (
*Spelobia ibrida Roháček, 1983 – TPH
Material. Kubus-larger, sweep-netting, 14.ix.2018, 1♀. Kubus-smaller, sweep-netting, 14.ix.2018, 1♂ (genit. prep.).
Comments. An uncommon phytosaprophagous species known from montane regions of Central and South Europe (
Spelobia luteilabris (Rondani, 1880) – TN
Material. Kurnoyatsu-2, dry habitat (Sphagnum fuscum), sweep-netting, 7.vi.2018, 1♂ (genit. prep.).
Comments. A polysaprophagous, habitat- and altitude-tolerant, originally Holarctic species. It is also known from some European peat bogs (
Spelobia parapusio (Dahl, 1909) – TN
Material. Kurnoyatsu-2, dry habitat (Sphagnum fuscum), yellow pan traps, 22–24.ix.2018, 1♂ (genit. prep.). Kubus-smaller, yellow pan traps, 12-14.ix.2018, 1♂. Tarskoe, sweep-netting, 11.ix.2018, 1♂.
Comments. An originally Palaearctic mycophagous species developing in sporocarps of various terrestrial macrofungi, mainly in woodland. It is parthenogenetic in Central and North Europe but bisexual in southern parts of the Palaearctic Region. Spelobia parapusio was described from 1♀ collected in Sphagnum on a raised bog in Germany (
Spelobia rufilabris (Stenhammar, 1855) – TN
Material. Kubus-smaller, sweep-netting (1), 12.ix.2018, 1♀; same bog, yellow pan traps, 12-14.ix.2018, 1♂ (genit. prep.).
Comments. A phytosaprophagous Palaearctic species living mainly in colder and humid montane forests but (less frequently) also occurring on peat bogs (
Spelobia talparum (Richards, 1927) – TN
Material. Kubus-smaller, sweep-netting (1), 12.ix.2018, 1♀ (genit. prep.).
Comments. A common microcavernicolous species, widespread in the Palaearctic Region. It is polysaprophagous and inhabits subterraneous nests and runs of mammals. There are several records from peat bogs in the Czech Republic (
Terrilimosina schmitzi (Duda, 1918) – TN
Material. Kubus-smaller, sweep-netting, 4.vi.2018, 1♂ (genit. prep.).
Comments. A widespread Holarctic species associated with forest litter in humid montane woodland. It was also ascertained in peat bogs in Wales (
A total of 38 species of Sphaeroceridae has been found in eight montane and submontane mires of the North Caucasus. Sphaeroceridae appeared to be rather diverse but not abundant in the insect fauna of these mires, as compared to members of many other dipteran families, which were much more common. Extensive sampling with seven main techniques provided only 119 specimens of Sphaeroceridae. Most Sphaeroceridae species (31) were represented only by 1–3 specimens; most species (32) were found only in one or two mires. Only one species, Spelobia clunipes, was relatively common and, at the same time, it was recorded from the highest number of mires (5 of 8). The highest species number (17) was recorded in the habitat-diverse Kurnoyatsu-2, the lowest (2–5), in the most uniform and largest fens of Ushtulu and Chifandzar and in a relatively uniform small mire Kurnoyatsu-3.
Because Sphaeroceridae are usually partly or wholly neglected in studies on insect communities of peat bogs (see Introduction), the above results obtained on the Caucasian mires can only be compared with those from more detailed investigations of this family in the Czech Republic (
In the Caucasian mires under study only one (and new) tyrphobiont species was found, Pullimosina (P.) turfosa sp. nov. In Europe, three other species have been formerly placed in this category: two circumpolar (hence Holarctic) Boreo-montane species of Limosininae, Pullimosina (Dahlimosina) dahli (Duda, 1918) and Spelobia pappi Roháček, 1983, and one species of Copromyzinae hitherto only known from raised bogs in the Czech Republic and Austria, Crumomyia tyrphophila Roháček, 1999 (see
The following ten species have been classified as tyrphophilous by
The vast majority (34 species) of Sphaeroceridae recorded from Caucasian mires belong to this category which is in agreement with the situation in Central and North European bogs. This group is mainly composed of common eurytopic coprophagous or saprophagous species, which are able to survive (and even prosper) in peat bogs if there is enough food supply for their larvae (animal excrement and carrion and decaying vegetation). Five of the eight mires under study can provide favorable conditions to many coprophagous species of Sphaeroceridae as their environs and essentially the bogs are used for grazing horses or cattle (see “Localities under study” for further details). Borborillus, Copromyza, Lotophila, Norrbomia, Chaetopodella, Coproica, Gonioneura, and Opalimosina species are typical examples of coprophages while Eulimosina, Leptocera, Minilimosina, Pullimosina, and most Spelobia and Terrilimosina species are phytosaprophagous or polysaprophagous. The herein recorded paludicolous Opacifrons, Pseudocollinella, and Rachispoda species are also eurytopic and can develop in the acidic mud in these mires, perhaps with the exception of Rachispoda hostica (Villeneuve, 1917) being obviously an occasional vagrant and hence belonging to the tyrphoxenous category. More interesting is the occurrence of microcavernicolous species living in burrows of small mammals, viz. Spelobia czizeki (Duda, 1918) and S. talparum (Richards, 1927). The mycophagous Spelobia parapusio (Dahl, 1909) is interesting because only males have been found although it is a parthenogenetic species at higher latitudes (
Wing reduction and the loss of or decreased ability to fly has been observed in many different families of Diptera (e.g., Tipulidae, Limoniidae, Chironomidae, Phoridae, Chloropidae, Ephydridae, Drosophilidae, etc.), primarily in species adapted to cold and open windy habitats, and especially in those living at high latitudes and altitudes (e.g., see
A.A. Przhiboro is grateful to Yu.A. Dunaeva (St. Petersburg, Russia) for her help in fieldwork. Both authors are very grateful to D.A. Philippov (Borok, Yaroslavl Province of Russia) for the identification of monocotyledones and comments on mire typology, to M.A. Boychuk (Petrozavodsk, Russia) for the identification of Sphagnum mosses, to P. Chandler (Melksham, England) for comments and English revision of the manuscript, and to the reviewers for helpful criticism. The research of Sphaeroceridae by J. Roháček was supported by the Ministry of Culture of the Czech Republic by institutional financing of long-term conceptual development of research institution (the Silesian Museum, MK000100595). The work of A. Przhiboro was conducted within the framework of State Assignment No. 122031100274-7, Ministry of Science and Higher Education of the Russian Federation.