The Biological Diversity of Fruit Flies (Diptera: Drosophilidae) in Russia: A Description of a Set of Own and Published Data and a Complete List of Species

Abstract

Drosophilidae is a relatively small family within Diptera. However, species of this family occupy a wide range of ecological niches and are frequently found in synanthropic habitats. Additionally, some species are known agricultural pests. The dataset is based on collections of Drosophilidae from eleven regions of Russia. The dataset was uploaded to the GBIF platform in 2024. Published sources specifying exact localities and collection dates were also used. The database includes records dating back to 1867, with the majority of specimens collected by the authors between 2001 and 2024. Collection methods included net sweeping and bait trapping. The dataset contains 2830 occurrence records, with a total of 51,006 specimens of Drosophilidae studied. It includes data on 108 species from two subfamilies, covering 49 regions of Russia. Considering additional published sources, 188 species of Drosophilidae are currently known from Russia, with a complete species list provided. Among the most abundant species in the dataset, 10 species are represented by more than 1000 specimens: Drosophila obscura, Scaptodrosophila rufifrons, Drosophila melanogaster, Drosophila phalerata, Drosophila transversa, Drosophila kuntzei, Drosophila histrio, Drosophila testacea, Phortica semivirgo, and Drosophila immigrans. Conversely, 39 species are represented by fewer than 10 specimens in the dataset. Regarding ecological groupings, the most dominant groups are xylosaprobionts (39 species, 40.6%) and mycophages (30 species, 31.3%). Notably, in 2017, the quarantine pest Drosophila suzukii was detected in the European part of Russia. The current knowledge of the Drosophilidae fauna in Russia remains insufficient. Of the 15 regions, only one or two localities are represented in the dataset. The distribution limits and range boundaries of many species remain unknown. Moreover, the local faunas of more than half of Russia’s regions remain unexplored.

1. Summary

The study of species diversity in animals and plants has long attracted the attention of taxonomists, ecologists, and other specialists [1,2]. In recent years, this issue has received increased attention due to the observed decline in population numbers and biodiversity worldwide [3,4]. A key question concerns the biodiversity of taxonomic groups. To facilitate information exchange, specialized databases have been developed, among which the Global Biodiversity Information Facility (GBIF) is one of the largest. The GBIF contains over 2 billion occurrence records of various organisms, and this number continues to grow [5,6].

Diptera is one of the most challenging insect orders to study. These insects are highly active, sometimes abundant, sometimes rare, and morphologically diverse, with a global distribution across all continents. Their vast range of forms and sizes has enabled them to occupy a wide variety of ecological niches, both in natural habitats and in anthropogenically altered landscapes [7,8,9,10].

Drosophilidae are small Diptera, typically yellow to dark brown in coloration, often with reddish eyes. This family is widely distributed across the globe, with more than 4600 known species worldwide and over 300 species recorded in the Palearctic region [11]. The biodiversity of fruit flies is particularly high in tropical regions and the southern Palearctic. In Brazil, 304 species have been recorded [12], while a single Afrotropical island, São Tomé in the Indian Ocean, harbors 80 species [13]. In the southern Palearctic, Bulgaria hosts 120 species [14].

In the northern Palearctic, the species diversity of Drosophilidae decreases significantly. For instance, Canada has 79 known species, with an estimated additional 20–25 species potentially present [15]. Finland has recorded 67 species [16], and Norway 47 species [17]. Previous estimates suggested that Russia hosts between 200 and 230 species [18]. However, more recent assessments based on targeted studies of this family estimate the Russian fauna at approximately 160 species [19].

Drosophilidae inhabit various natural biotopes, often found on tree bark near exuding sap, on flowers, and on ripe fungi [20,21]. Many species are synanthropic and have spread into human dwellings. In synanthropic habitats, they are commonly found near fruits, vegetables, garbage heaps, and other organic substrates [22,23,24]. Drosophilidae are particularly attracted to fermenting substrates [25,26].

Larvae develop in various decaying substrates, feeding on microorganisms that thrive in these environments. They inhabit rotting berries, fungi, fruits, wood, and tree bark with exuding sap. Some species mine the stems and leaves of various agricultural plants, causing economic damage [27,28,29].

The objective of our study was to compile a database based on research conducted from 2019 to 2024, supplemented by reliable published data with precise locality names and/or geographic coordinates of Drosophilidae species occurrences. The study aimed to achieve the following: (1) Determine the number of Drosophilidae species in the fauna of Russia; (2) Identify abundant and rare species; (3) Classify species into ecological groups; (4) Detect invasive and potentially harmful pest species.

2. Data Description

2.1. Dataset Description

In the described dataset, each recorded observation includes key information such as location (latitude/longitude), date of observation, observer’s name, and identifier’s name. If the observation has been previously published, a reference is provided. Coordinates were determined on-site using a GPS device or post-study through Google Maps (

Table 1. Description of the data in the dataset [30].

Column Label	Column Description
occurrenceID	An identifier for the occurrence (as opposed to a particular digital record of the occurrence)
basisOfRecord	The specific nature of the data record: HumanObservation
scientificNam	The full scientific name including the genus name and the lowest level of taxonomic rank with the authority
Kingdom	The full scientific name of the kingdom in which the taxon is classified
decimalLatitude	The geographic latitude of the location in decimal degrees
geodeticDatum	The ellipsoid, geodetic datum, or spatial reference system (SRS) upon which the geographic coordinates given in decimalLatitude and decimalLongitude is based
Country	The name of the country in which the location occurs
countryCode	The standard code for the country in which the location occurs
individualCount	The number of individuals present at the time of the occurrence
eventDate	The date when material from the trap was collected or the range of dates during which the trap collected material
Year	The integer year in which the event occurred
Month	The ordinal month in which the event occurred
Day	The integer day of the month on which the event occurred
recordedBy	A person, group, or organization responsible for recording the original occurrence
identifiedBy	A list of names of people who assigned the taxon to the subject
bibliographicCitation	A bibliographic reference for the resource

).

2.2. Biodiversity of Fruit Flies

The dataset includes records of 108 Drosophilidae species from two subfamilies (Steganinae and Drosophilinae) across 49 regions of Russia (

Table 2. Species diversity of Drosophilidae in Russia (dataset).

Species	Specimens	Locality
Steganinae
Amiota albilabris (Roth, 1860)	88	26
Amiota alboguttata (Wahlberg, 1839)	89	23
Amiota flagellata Okada, 1971	4	1
Amiota rufescens (Oldenberg, 1914)	41	16
Amiota subtusradiata (Duda, 1934)	54	20
Cacoxenus vlasovi (Duda, 1934)	1	1
Gitona distigma Meigen, 1830	144	36
Leucophenga maculata (Dufour, 1839)	53	14
Leucophenga quadripunctata (Meijere, 1908)	2	1
Leucophenga quinquemaculata Strobl, 1893	717	40
Leucophenga quinquemaculipennis Okada, 1956	9	2
Leucophenga sorii Kang, Lee & Bhang 1965	3	3
Leucophenga stackelbergi Duda, 1934	10	4
Phortica chi (Toda & Sidorenko, 1996)	17	6
Phortica conifera (Okada, 1977)	60	5
Phortica iota (Toda & Sidorenko, 1996)	141	10
Phortica semivirgo (Máca, 1977)	1327	33
Phortica variegata (Fallén, 1823)	9	6
Stegana baechlii Laštovka & Máca, 1982	1	1
Stegana coleoptrata (Scopoli, 1763)	11	10
Stegana furta (Linnaeus, 1767)	16	7
Stegana hypoleuca Meigen, 1830	10	5
Stegana longifibula Takada, 1968	7	4
Stegana masanoritodai Okada & Sidorenko, 1992	1	1
Stegana mehadiae Duda, 1924	3	2
Stegana sibirica (Duda, 1934)	14	1
Stegana similis Laštovka & Máca, 1982	2	2
Drosophilinae
Chymomyza amoena (Loew, 1862)	519	21
Chymomyza caudatula Oldenberg, 1914	40	19
Chymomyza costata (Zetterstedt, 1838)	167	23
Chymomyza distincta (Egger, 1862)	33	6
Chymomyza fuscimana (Zetterstedt, 1838)	89	21
Drosophila alpina Burla, 1948	188	12
Drosophila altukhovi Imasheva, Lazebny, Cariou, David & Tsacas, 1994	1	1
Drosophila ambigua Pomini, 1940	1	1
Drosophila angularis Okada, 1956	17	4
Drosophila bifasciata Pomini, 1940	355	26
Drosophila busckii Coquillett, 1901	211	17
Drosophila ezoana Takada & Okada, 1957	133	7
Drosophila funebris (Fabricius, 1787)	956	29
Drosophila helvetica Burla, 1948	38	7
Drosophila histrio Meigen, 1830	2426	43
Drosophila hydei Sturtevant, 1921	147	18
Drosophila imaii Moriwaki & Okada, 1967	10	1
Drosophila immigrans Sturtevant, 1921	1226	29
Drosophila ingrica Hackman, 1957	4	3
Drosophila kuntzei Duda, 1924	2978	48
Drosophila limbata Roser, 1840	24	10
Drosophila littoralis Meigen, 1830	6	6
Drosophila lummei Hackman, 1972	1	1
Drosophila melanogaster Meigen, 1830	4445	46
Drosophila mercatorum Patterson & Wheeler, 1942	35	2
Drosophila montana Stone, Griffen & Patterson, 1941	42	5
Drosophila nigromaculata Kikkawa et Peng, 1938	22	7
Drosophila nigrosparsa Strobl, 1898	11	1
Drosophila obscura Fallén, 1823	13,480	49
Drosophila parakuntzei Okada, 1973	2	2
Drosophila phalerata Meigen, 1830	4132	50
Drosophila picta Zetterstedt, 1847	2	2
Drosophila rellima Wheeler, 1960	18	2
Drosophila repleta Wollaston, 1858	41	7
Drosophila simulans Sturtevant, 1919	6	1
Drosophila subobscura Collin, 1936	311	23
Drosophila subsilvestris Hardy & Kaneshiro, 1968	362	21
Drosophila suzukii (Matsumura, 1931)	12	5
Drosophila testacea Roser, 1840	2839	56
Drosophila transversa Fallén, 1823	3891	79
Drosophila tristis Fallén, 1823	16	6
Drosophila unispina Okada, 1956	20	3
Drosophila virilis Sturtevant, 1916	6	3
Hirtodrosophila baikalensis Watabe, Toda & Sidorenko, 1996	7	3
Hirtodrosophila cameraria (Haliday, 1833)	13	6
Hirtodrosophila confusa (Staeger, 1844)	165	30
Hirtodrosophila histrioides (Okada & Kurokawa, 1957)	31	3
Hirtodrosophila lundstroemi (Duda, 1935)	1	1
Hirtodrosophila oldenbergi (Duda, 1924)	4	2
Hirtodrosophila sexvittata (Okada, 1956)	56	2
Hirtodrosophila shaitanensis Sidorenko, 1996	1	1
Hirtodrosophila subarctica (Hackman, 1969)	317	22
Hirtodrosophila toyohiokadai (Sidorenko, 1990)	54	10
Hirtodrosophila trivittata (Strobl, 1893)	59	20
Lordiphosa acuminata (Collin, 1952)	3	2
Lordiphosa andalusiaca (Strobl, 1906)	1	1
Lordiphosa fenestrarum (Fallén, 1823)	17	11
Lordiphosa miki (Duda, 1924)	3	1
Lordiphosa mommai (Takada & Okada, 1960)	560	1
Lordiphosa nigricolor (Strobl, 1898)	11	6
Microdrosophila congesta (Zetterstedt, 1847)	2	2
Microdrosophila cristata Okada, 1960	1	1
Mycodrosophila celesta Sidorenko, 1992	7	2
Mycodrosophila poecilogastra (Loew, 1874)	59	7
Mycodrosophila shikokuana Okada, 1956	8	1
Mycodrosophila takachihonis Okada, 1956	1	1
Nesiodrosophila magnidentata (Lee, 1964)	1	1
Scaptodrosophila coracina (Kikkawa & Peng, 1938)	54	2
Scaptodrosophila deflexa (Duda, 1924)	92	2
Scaptodrosophila rufifrons (Loew, 1873)	6374	39
Scaptomyza carinata Okada, 1973	216	6
Scaptomyza consimilis Hackman, 1955	27	11
Scaptomyza flava (Fallén, 1823)	50	20
Scaptomyza graminum (Fallén, 1823)	59	22
Scaptomyza okadai Hackman, 1959	13	7
Scaptomyza pallida (Zetterstedt, 1847)	538	43
Scaptomyza taigensis Sidorenko & Toda, 1996	8	2
Scaptomyza teinoptera Hackman, 1955	2	1
Scaptomyza unipunctum (Zetterstedt, 1847)	13	9
Scaptomyza yakutica Sidorenko & Toda, 1996	80	2
Zaprionus flavofasciatus (Takada, Beppu & Toda, 1979)	1	1
Total	51,006

). The total number of occurrences recorded is 2830, with 51,006 specimens documented in the dataset.

In the identification guide for the European part of Russia, Drosophilidae are estimated to include 44–54 species [31]. A more recent publication on the Russian Far East reports that over 160 species inhabit Russia [19]. We have attempted to expand the list of Drosophilidae species by including those not recorded in our dataset. Appendix A provides a list of these species, compiled based on published sources [19,32].

Considering all available data, the current fauna of Russia comprises 188 Drosophilidae species. The Russian Far East remains the most thoroughly studied region, exhibiting significant species diversity. However, the European part of Russia has also been increasingly studied in recent years. It is possible that, with continued intensive research, the number of documented species may exceed 200.

Comparing the Drosophilidae fauna of Russia with other countries presents a challenge due to Russia’s vast territory, unparalleled by other nations. However, for reference, some well-studied countries report the following numbers of species: 79 in the Czech Republic, 71 in Slovakia, and 120 in Bulgaria [14,33].

Among the Drosophilidae recorded in the dataset, 10 species stand out, each with more than 1000 registered specimens. At the same time, these species also have a significant number of documented localities. However, for three species (Gitona distigma, Leucophenga quinquemaculata, and Scaptomyza pallida), despite having lower overall specimen counts, the number of recorded localities is notably high (Figure 1).

Drosophila obscura is a typical xylosaprobiont species, with larvae primarily living in decaying tissues under the bark and in fermenting tree sap of various deciduous trees. The larvae of Phortica semivirgo and Scaptodrosophila rufifrons are also xylosaprobionts, sharing the same habitats as Drosophila obscura. In contrast, the larvae of Drosophila histrio, Drosophila kuntzei, Drosophila phalerata, Drosophila testacea, and Drosophila transversa are mycetobionts, living in various fungi, predominantly in live fungi, but sometimes in decaying ones.

Among the most abundant Drosophilidae species found in Russia, Drosophila immigrans and Drosophila melanogaster are synanthropic species, closely associated with human activity. These species inhabit and breed in areas with fermenting and decaying fruits, vegetables, wine, beer, and juices. Although they are primarily found in human-influenced environments, these species may occasionally occur in small numbers in wild biotopes, likely due to migration or wind dispersal.

Drosophilidae are unevenly distributed in forest biotopes, with most species preferring the lower tiers of the forest. In boreal and cool temperate forests in Russia, canopy-dwelling Drosophilidae species are mostly sap and decayed bark feeders. The largest group of Drosophilidae inhabiting the lower forest tiers consists primarily of mycetobiont species and some xylosaprobiont species, including the most abundant Drosophila obscura, Phortica semivirgo, and Scaptodrosophila rufifrons [34]. Drosophilidae are also distributed unevenly across different types of forests in European Russia, being most numerous in birch forests [35].

A total of 39 species were recorded in the database, with each species having 10 or fewer specimens. Thirteen species were represented by only one specimen (Cacoxenus vlasovi, Drosophila altukhovi, Drosophila ambigua, Drosophila lummei, Hirtodrosophila lundstroemi, Lordiphosa andalusiaca, Microdrosophila cristata, Mycodrosophila takachihonis, Nesiodrosophila magnidentata, Stegana masanoritodai, Stegana baechlii, Zaprionus flavofasciatus, Hirtodrosophila shaitanensis). These species were found in regions such as Krasnoyarsk Krai, Khabarovsk Krai, Primorsky Krai, the Republic of Dagestan, Saint Petersburg, Sakhalin Oblast, and the Republic of Altai.

Five species were found in two specimens each (Leucophenga quadripunctata, Microdrosophila congesta, Drosophila parakuntzei, Drosophila picta, Stegana similis), and these species are distributed across only seven regions. Four species (Leucophenga sorii, Lordiphosa acuminata, Stegana mehadiae, Lordiphosa miki) were represented by three specimens each, found in five regions.

In total, for the 39 species in the dataset, only 10 or fewer specimens are known from six or fewer localities. This means that 36.1% of the species are recorded from one, two, or three regions within Russia. This highlights the limited study of much of Russia’s territory, with the exception of the Far East and, to some extent, parts of the Caucasus. Of the 15 regions, only one or two localities are represented in the dataset. In fact, the fauna of half of Russia’s regions, including the central, western, Volga, and Siberian areas, is poorly documented.

Based on the dataset, we have prepared an analysis of the ecological groups of Drosophilidae according to their larval breeding sites (Figure 2).

Information was obtained for 96 species, with the main groups being xylosaprobionts (39 species, 40.6%) and mycophages (30 species, 31.3%). It has been previously noted [34,36] that in the temperate zone, most species of this family belong to the aforementioned groups. Specifically, eight out of the ten most numerous species are classified as xylosaprobionts (Drosophila obscura, Scaptodrosophila rufifrons, Drosophila testacea, Phortica semivirgo) and mycophages (Drosophila phalerata, Drosophila transversa, Drosophila kuntzei, Drosophila histrio). Eight species are listed as saprophages and phytosaprophages. One species, Cacoxenus vlasovi, is classified as a kleptoparasite, inhabiting the nests of Osmia and Chalicodoma (Hymenoptera) [37].

One of the most well-known invasive Drosophilidae species is Drosophila suzukii. This fruit pest is a polyphage that can pierce the skins of soft fruits while laying its eggs. In Europe, Drosophila suzukii was first discovered in Spain and Italy in 2008, and soon after, it spread to many other countries (France, Austria, Germany, Belgium, Croatia, Slovenia, Ireland, UK, and Switzerland) [38,39]. In Russia, the first findings of the species were recorded in the Far East and the Caucasus. In the Krasnodar region, the species has already spread actively, causing damage to grapes, figs, and other fruit crops [40,41,42].

Two other species also cause certain damage to agricultural crops. Scaptomyza flava is a polyphage and has been found on Brassicaceae, Asteraceae, Capparidaceae, Resedaceae, Tropaeolaceae, and Viciaceae in European and other countries. It frequently damages plants such as Brassica rapa, Brassica campestris, Brassica juncea, Eruca vesicaria, Cardamine debile, Cardamine flexuosa, Cardamine hirsuta, Rorippa nasturtium-aquaticum, Coronopus didymus, Raphanus raphanistrum, Raphanus maritimus, and other crops [43]. Scaptomyza graminum mines the leaves of Caryophyllales, Amaranthaceae, Portulaceae, Fabaceae, causing damage to crops such as Beta vulgaris, Brassica campestris, Brassica oleracea, Brassica rapa, and others [44,45].

Also included in the group of pests are synanthropic Drosophilidae species (Drosophila melanogaster, Drosophila simulans, Drosophila hydei, Drosophila repleta, Drosophila immigrans, Drosophila busckii, Drosophila virilis, Drosophila funebris), which reproduce in large numbers on fruits, vegetables, juices, beer, and wine, contaminating the products with their adults and larvae [46,47].

3. Methods

3.1. Study Area

Descriptions of the nature and territory of Russia can be found in many publications. The authors’ own research was conducted within the boundaries of the Volga Upland and the Oka–Don Lowland. The majority of this large macroregion is located in the mixed forest zone, with broadleaf forests and forest steppe in the south and taiga forests in the northeast. The largest rivers are the Don (and its numerous tributaries), which flows from the west and southwest, and the Oka, which flows from the northern side of the macroregion. To the east, the region is bounded by the Volga River (Figure 3).

Figure 3 shows the locations from which data were obtained for the dataset. Due to the lack of data from many other regions of Russia, significant gaps are visible on the map (as noted above).

3.2. Study Material

The material for the dataset was collected from Drosophilidae specimens across 49 regions of Russia. In addition, reliable publications were used, which provide trust in the original data sources. The earliest Drosophilidae specimen dates back to 1867. However, the majority of the material (2303 occurrences, 81.4%) was collected between 2001 and 2024. The dataset did not include materials from the publications of vs. Sidorenko, as virtually all of his results were used in two well-known papers [19,32].

In our own research, traditional and various methods were employed. Primarily, these were collections using nets, as well as trapping with various baits [48]. After capture, Drosophilidae were preserved in a 70% ethanol solution for subsequent identification. Identification was performed at the Laboratory of Evolutionary Developmental Genetics of the N.K. Koltsov Institute of Developmental Biology, RAS, Moscow, Russia. A Micromed MS-2-Zoom microscope (Moscow, Russia) was used to determine the drosophilids. The identification was carried out by N.G. Gornostaev using drosophilid identification keys [19,49]. The systematics of the Drosophilidae was interpreted according to Grimaldi [50].