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65р-ISSN 2617-6157 е-ISSN 2617-6165 GEO&BIO • 2020 • том 19
e current composition of the family Gobiidae Cuvier,
1816 (Actinopterygii) inwaters ofUkraine with comments
on species distributions
Leonid Manilo https://orcid.org/0000-0002-7143-9470
National Museum of Natural History, NAS of Ukraine (Kyiv, Ukraine)
e current composition of the family Gobiidae Cuvier, 1816 (Actinopterygii) in waters of Ukraine with
comments on species distributions. — L. Manilo. — e present paper is devoted to a literature-based ret-
rospective analysis of changes in the species composition of the family Gobiidae of waters of Ukraine since
the 1930s. Keys and checklists published earlier cover the sh fauna of separate regions of Ukraine, either
only of river basins or of coastal waters of the Black Sea and the Sea of Azov. e rst checklist of shes of
Ukraine was published by academician A. M. Nikolsky in 1930 including both freshwater and marine spe-
cies known in that time. In that work, the family Gobiidae includes 23 species of 7 genera. In the paper by D.
A. Tretyakov (1947), 23 species of 10 genera of true gobies are listed for waters of Ukraine with some chang-
es in the species composition. In one of the volumes of the series “Fauna of Ukraine. Fishes” devoted to true
gobies (Smirnov 1986), data on 25 species of 11 genera are presented. According to the data by Yu. Movchan
(2011), the composition of the family largely expanded comprising 33 species of 13 genera, including 5
marine species that were recorded in that time in coastal waters of the Crimea as well as 1 newly described
species from the Chorna River. Later, ve more marine species, known earlier only in the Mediterranean Ba-
sin, were found in waters of Ukraine, as well as another species in the Sea of Azov. It was established that the
current sh fauna of the family Gobiidae of Ukraine includes 38 species of 17 genera, according to the lit-
erature and data from the sh collection of the National Museum of Natural History NAS of Ukraine (Kyiv).
e geographic range and distribution of each species in waters of Ukraine are presented. e fauna of true
gobies of Ukraine is composed of species of the Ponto-Caspian (52.6 %) and Atlantic-Mediterranean (47.4
%) faunal complexes. Endemics (4species) are highlighted along with recent intruders of the Black Sea that
have appeared here since the early 2000s (8 species); the time of appearance for 2 species remains unknown.
Key w ords: Gobiidae, genus, species, distribution, Ukraine, Black Sea, Sea of Azov, faunal complex,
endemic species.
Introduction
True gobies (Gobiidae) are one of the largest family of the order Gobiiformes that includes 1359
species of 189 genera (Nelson et al. 2016) distributed in tropical and temperate latitudes from the
intertidal zone to a depth of 200 m. True gobies are important components of sh diversity having a
signicant role in trophic chains of water ecosystems. In the sh fauna of Ukraine, the family of true
gobies is the second largest by the number of species aer the cyprinids (Cyprinidae).
Many of the earlier published checklists and keys to shes cover mainly the whole sh fauna of
separate regions (Ostroumov 1896; Sushkin & Beling 1923; etc.), while others deal only with freshwa-
ter sh faunas (Markevych & Korotkyy 1954), species composition of shes of separate river basins,
regions, or coastal waters of Ukraine (Vladykov 1926; Kolyushev 1949; Lyashenko 1952; Manilo 2014;
etc.). erefore, these publications cannot present the entire family of true gobies.
One of the rst publications with a description of the sh fauna of Ukraine was the work by aca-
demician A. M. Nikolsky (1930), in which the author considers not only freshwater, but also marine
species of coastal waters of Ukraine in the Black Sea and the Sea of Azov. In this key, the family
Gobiidae is represented by 23species of 7 genera, such as Gobiosoma, Aphia, Pomatoschistus, Gobius,
Proterorhinus, Benthophilus and Benthophiloides.
e systematics of true gobies started to develop intensely aer the studies of the lateral line sys-
tem in dierent genera and species by B. S. Ilin (1927, 1949). In the “Key to Cyclostomes and Fishes of
Correspondence to: Leonid Manilo; National Museum of Natural History, NAS of Ukraine; Bohdan Khmelnytsky
St. 15, Kyiv, 01030 Ukraine; е-mail: leonid.manilo@gmail.com; orcid: 0000-0002-7143-9470
2020 • vol. 19, pp 65–84
https://doi.org/
GEO&BIO
66 GEO&BIO • 2020 • vol. 19 р-ISSN 2617-6157 е-ISSN 2617-6165
the Ukrainian SSR” by D. K. Tretyakov (1947), 23 species of 10 genera of true gobies are presented for
waters of Ukraine (three genera are added: Knipowitschia, Zostericola, Bubyr along with modications
in the species composition), and gobies are rst considered among other local literature sources as a
separate order Gobiiformes.
In the following years, many papers and monographs were published devoted to dierent aspects
of systematics, biology, and distribution of gobies. In 1980–1988, the series “Fauna of Ukraine. Fishes”
was published (issues 1–5 in six books), including an issue dedicated to gobies (Smirnov 1986). In that
publication, the author considers 25 species of 11 genera in the composition of the family Gobiidae.
Yu. V. Movchan (2011), based on a survey and analysis of novel ichthyological data, generalised
the materials of numerous sources on shes of waters of Ukraine, including gobies. According to
this publi cation, the composition of Gobiidae signicantly expanded up to 33 species of 12 genera,
inclu ding marine species (Pomatoschistus bathi, Millerigobius macrocephalus, Gobius cruentatus, and
G. xanthocephalus) that were discovered in coastal waters of the Crimea by scientists of the Institute of
Biology of Southern Seas, NAS of Ukraine (Boltachev et al. 2009a, 2010; Boltachev & Karpova 2010).
Additionally, a new species Proterorhinus tataricus was described from the Chorna River (Freyhof &
Naseka 2007).
Later, several other species were discovered in coastal waters of the Crimea previously known
to be occurring only in the Mediterranean Basin: Chromogobius quadrivittatus (Kovtun 2013),
Gammogobius steinitzi (Kovtun & Manilo 2013), as well as Benthophilus magistri from the Sea of
Azov (Manilo 2011a). ese species were included into L. G. Manilo’s monograph, and the number of
species and genera increased to 34 and 17, respectively (Manilo 2014).
Aerwards, a number of Mediterranean species were discovered in coves of Sevastopol and under-
water caves of Cape Tarkhankut: Chromogobius zebratus, Gobius couchi, and Zebrus zebrus (Kovtun &
Karpova 2014; Karpova & Boltachev 2018).
However, there are no publications dealing with the complete current composition of the family
of waters of Ukraine, including both freshwater and marine species. erefore, the aim of the present
paper is to generalise knowledge and to determine the current composition of the family Gobiidae
along with the distribution of species.
Material and Methods
e study is based on results of a retrospective analysis of literature on the composition and dis-
tribution of shes of the family Gobiidae occurring in waters of Ukraine. e systematics and no-
menclature of true gobies are presented based on their validity according to modern ichthyological
research (Froese & Pauly 2019).
Based on recent data of phylogenetic and biogeographic analyses (Neilson & Stepien 2009), the
expansion of the taxonomic volume of subfamily Benthophilinae (based on the tribe Benthophilini
Beling et Iljin, 1927) by inclusion of the genus Neogobius and other close genera is substantiated.
is subfamily is divided into three tribes: Benthophilini, Neogobiini, and Ponticolini. e tribe
Ponticolini, in addition to the genera Mesogobius and Proterorhinus, also includes Babka and Ponticola
as independent genera. us, the system of the family of true gobies is presented in this paper accord-
ing to these data.
e unit of zoogegographic analysis is a faunal complex, i.e. a group a species connected by com-
mon geographic origin (Nikolsky 1947). Due to the complex geological past of the Azov–Black Sea
Basin, its sh fauna, including the family Gobiidae, has a multifarious origin and includes repre-
sentatives of dierent faunal complexes such as Boreal-Atlantic, Mediterranean, and Ponto-Caspian
(Zenkevich 1963). According to more recent views (Zaitsev 1998), the ichthyofauna of the Black
Sea was ultimately formed aer the last connection between the Black Sea and the Mediterranean
Sea. Species of the Boreal-Atlantic complex had already been present in the compostion of the
Mediterranean complex before the Dardanelles appeared, thus this complex is considered in this
paper as Atlantic-Mediterranean.
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All species of the Ponto-Caspian complex are Ponto-Caspian endemics by their origin. In this pa-
per, endemics are considered based on their current distribution, i.e. a species is considered endemic
if its distribution is restricted to the Black Sea and Sea of Azov or has an even narrover range (e.g. Pr.
tataricus).
Among the representatives of the Atlantic-Mediterranean complex, recent invaders that appeared
in the Azov-Black Sea Basin in the last few decades are highlighted.
Systematic Part
Order Gobiiformes
Family Gobiidae
Subfamily Benthophilinae
Tribe Benthophilini Neilson et Stepien, 2009
Genus Benthophiloides Beling et Iljin, 1927
e genus includes two species distributed in estuaries and rivers of the Black and Caspian Seas.
One species occurs in waters of Ukraine.
Benthophiloides brauneri Beling et Iljin, 1927
A species of the Ponto-Caspian faunal complex. e only representative of the genus in the fauna
of Ukraine. Inhabits estuaries, bays, river mouths, reservoirs, and coastal waters of the northwest part
of the Black Sea and the Sea of Azov. In Ukraine, it was recorded in the middle and upper sections
of the Dnipro Estuary (Snigirov 2014), in Odesa Bay (Cape Velyky Fontan), in the Bug Estuary from
Nova Odesa to Mykolaiv, in the course of the Dnipro from Kherson to Kakhovka, in Tendra Bay, and
in the upper sections of the Dnipro (Novitsky et al. 2008; Stepanok & Hubanov 2018; Tkachenko
2018) and Kaniv Reservoirs (Manilo et al. 2013b). One record of the species is also known from Tatar
cove of Kazantip Bay, Sea of Azov (Boltachev et al. 2009b).
Genus Benthophilus Eichwald, 1831
According to data from dierent sources, the genus includes 18–20 species (Boldyrev & Bogutskaya
2007), most of which inhabit the Caspian Sea. In waters of Ukraine, three species occur (Manilo
2011). Some researchers (Shandikov & Goncharov 2008) suggest the presence of another species
Benthophilus durrelli Boldyrev et Bogutskaya, 2004 in the Siversky Donets River.
Benthophilus magistri Iljin, 1927
A species of the Ponto-Caspian faunal complex. Endemic to the Sea of Azov, where inhabits its
eastern part from Taganrog Bay (Mius and Yeya Estuaries) to Temryuk Bay (Akhtanizovskiy Estuary).
e species is absent in the western part of the Sea of Azov and in Lake Syvash. Occurs in Taganrog
Bay from its mouth between Bilosaraiska and Dovha Spits to the mouth of the Don River. In waters
of Ukraine, the species is recorded in the middle section of Taganrog Bay near Kryva Spit (Manilo
2011a, 2014).
Benthophilus nudus Berg, 1898
A species of the Ponto-Caspian faunal complex. Distributed in the northwest part of the Black
Sea and in adjacent estuaries, lakes, and rivers. In waters of Ukraine, the species occurs near Zmiinyi
Island, in the mouth of the Danube River and adjacent lakes, in the Sasyk Estuary (Demchenko &
Smirnov 2009), in coastal waters of the Danube–Dnister interuve, in the Budatsky, Dnister, Tiligul
(until 1990) (Shekk 2004), Berezansky, and Dnipro–Bug Estuaries, in the lower section of the Dnister
River to Bendery city (Berg 1949), in the Southern Bug River to Oleksandrivka, in the Dnipro River
(including its tributaries Samara (Novitsky et al. 2006) and Ingulets (Ambroz 1956)), in Kaniv
Reservoir (Goroshko et al. 1989), in the mouth of the Desna River and near Kyiv (Pinchuk et al.
1985), as well as in Tendra Bay.
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Benthophilus stellatus (Sauvage, 1874)
A species of the Ponto-Caspian faunal complex. Endemic to the Sea of Azov. According to litera-
ture data, the distribution range of the species covers the Sea of Azov to the Kerch Peninsula, also
including Lake Syvash (Boldyrev & Bogutskaya 2007). According to some researchers, as a result of
accidental introduction the species appeared in the Volga River’s basin where it dispersed through
several waterbodies and now occurs in reservoirs of the Don and Volga Rivers (Vasileva 2007). In
waters of Ukraine, the species occurs in the south-western part of the Sea of Azov to Cape Kazantip,
as well as along the northern coast from the Utlyuk Estuary to Kryva Spit, including the Molochnyi
Estuary (Demchenko et al. 2005; Diripasko et al. 2011).
Genus Caspiosoma Iljin, 1927
A monotypic genus.
Caspiosoma caspium (Kessler, 1877)
A species of the Ponto-Caspian faunal complex. Its geographic range covers the basins of the Black
Sea, Caspian Sea, and the Sea of Azov (Svetovidov 1964; Luzhnyak & Korneev 2006). In waters of
Ukraine, the species occurs in estuaries of the northwest part of the Black Sea, particularly in the
Sasyk (Moshu 2006), Berezansky, Dnipro–Bug, and Dnister Estuaries (Strautman & Pinchuk 1972),
as well as from the Dnipro Delta to Kakhovka and in the Southern Bug near Mykolaiv. Not recorded
in waterbodies of Ukraine in the last few years.
Tribe Neogobiini Neilson et Stepien, 2009
Genus Neogobius Iljin, 1927
e genus comprises four species that inhabit the basins of the Black Sea, Sea of Azov, and the
Caspian Sea. Two species occur in waters of Ukraine.
Neogobius uviatilis (Pallas, 1814)
A species of the Ponto-Caspian faunal complex. Widely distributed in the basin of the Black Sea
(in the Danube to Austria (Ahnelt et al. 1998; Jurajda et al. 2005; Szalóky et al. 2015), in the Dnister
to its upper course, in the Southern Bug and Dnipro and their tributaries) and of the Sea of Azov (riv-
ers of the Azov Region and the Don with its tributaries). Records of the species are also known from
the Aegean (Zogaris & Apostolou 2011) and Marmara Seas (Sari et al. 2006). rough the system
of channels, the species penetrated the basin of the Baltic Sea (Danilkiewicz 1998; Rakauskas et al.
2018) and within the net of European freshwaters reached the Netherlands (Van Kessel et al. 2009).
In waters of Ukraine, the species occurs in coastal waters from the Danube Delta in the west to Kryva
Spit, Sea of Azov in the east, except for Zmiinyi Island and saline lakes Shagany, Alibei, and Burnas
(Manilo 2011b). Occurs practically in all rivers of Ukraine, including their middle and upper courses
(Smirnov 1986).
Neogobius melanostomus (Pallas, 1814)
A species of the Ponto-Caspian faunal complex. Inhabits the basin of the Black Sea, Sea of Azov,
and Caspian Sea (Svetovidov 1964), as well as of the Marmara and Aegean Seas (Eryilmaz 2002). In
the late 1950s, it was accidently introduced in the Aral Sea and occurred there until the 1980s, but by
now disappeared (Bogutskaya et al. 2004). e species got into the basin of the Baltic Sea with ballast
waters (Skóra & Rzeznik 2001). In recent years, it was recorded in the basin of the North Sea (Manné
et al. 2013). e species was accidently brought to North America, where it became naturalised and
widely distributed in the Great Lakes and rivers that ow into them (Jude 2001; Stepien & Tumeo
2006; Kornis et al. 2012). In the Black Sea and Sea of Azov, the species occurs along the shores. In
Ukraine, it inhabits along the entire coast of the Black Sea and Sea of Azov, in bays and estuaries ex-
cept for the saline lakes Shagany, Alibei, and Burnas (Manilo 2011b), as well as in deltas of all rivers.
e species reaches the middle and upper courses of the Danube, Dnister, Southern Bug, Dnipro, and
of rivers of the Azov Basin (Smirnov 1986; Movchan et al. 2003).
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Tribe Ponticolini Neilson et Stepien, 2009
Genus Babka Iljin, 1927
A monotypic genus.
Babka gymnotrachelus (Kessler, 1857)
A species of the Ponto-Caspian faunal complex. Inhabits desalinated estuaries, bays, coves, coastal
lakes, and rivers of the basin of the Black Sea, Sea of Azov, Caspian Sea, and Sea of Marmara. It pen-
etrated the basin of the Baltic Sea from the Dnipro through the system of canals (Danilkiewicz 1996;
Kottelat & Freyhof 2007), although the route of dispersal according to some other sources is un-
known (Grabowska 2005; Grabowska & Grabowski 2005). rough the Rhine–Main–Danube Canal,
it dispersed to Germany along the Danube (Roche et al. 2013). In waters of Ukraine, the species
was recorded in lakes of the Danube Delta area (Yalpug, Kagul, Katlabug, etc.), in the Sasyk Estuary
(Smirnov & Tkachenko 2007), in the Dnister from its mouth to the upper course and in its tributar-
ies Zbruch, Zhvanchyk, Strypa, Bystrytsia, Murafa, and Strviazh (Movchan et al. 2003; Grabowski et
al. 2016). It also occurs in the Southern Bug from its mouth to Ladyzhyn Reservoir, in the Dnipro
to Belorus, in the upper section of the Berezansky, Dnipro–Bug (Pinchuk 1977), and Tiligul (in pe-
riods of its desalination) estuaries (Shekk 2004), as well as in Yahorlyk, Dzharylhach, and Tendra
Bays (Manilo 2009a). In the basin of the Sea of Azov, the species occurs in the desalinated zone of
the northern part of the sea and in rivers of the Northern Azov Region (Smirnov 1986). It was also
recorded in reservoirs and lakes of eastern Crimea (Karpova 2009).
Genus Mesogobius Bleeker, 1874
e genus includes three species that inhabit marine, brackish, and freshwaters of the Azov–Black
Sea Basin and the Caspian Sea. One species occurs in waters of Ukraine.
Mesogobius batrachocephalus (Pallas, 1814)
A species of the Ponto-Caspian faunal complex. Its geographic range covers the Azov–Black Sea
Basin, the Bosporus, and the Sea of Marmara (Smirnov 1986). In waters of Ukraine, the species oc-
curs along the coasts, including Zmiinyi Island (Snigiryov 2008), Crimea (Boltachev 2003; Shaganov
2009), Kerch Strait (Shaganov & Milovanov 2005), western and northern parts of the Sea of Azov to
Taganrog Bay (Demchenko et al. 2005). It was also recorded in the lower course of the Danube, in
the middle course of the Dnister (Romanesku 2012) and the Southern Bug (to Oleksandrivka) and its
tributary Mertvovod, as well as in the Dnipro to Kyiv (Kakhovka, Dnipro, Kremenchug, and Kaniv
Reservoirs) (Smirnov 1986; Bulakhov et al. 2008) and in the Obitochna and Berda rivers. e species
was not found in saline lakes of the Danube–Dnister interuve (Shagany, Alibei, and Burnas), nor in
the Khadzhibey Estuary (Manilo 2009a).
Genus Ponticola Iljin, 1927
Initially, the genus Ponticola was described as a subgenus of the genus Gobius Iljin, 1927. In modern
systematics of the family, it is accepted as a valid genus, which currently includes 14Ponto-Caspian
species that inhabit the basins of the Black Sea, Sea of Azov, and Caspian Sea. In waters of Ukraine,
six species occur.
Ponticola cephalargoides (Pinchuk, 1976)
A species of the Ponto-Caspian faunal complex. Distributed in the western and north-western
parts of the Black Sea from the coast of Bulgaria and Romania to Cape Tarkhankut (western popula-
tion), in waters of the Kerch Peninsula from Cape Opuk in the Black Sea to Cape Kazantip in the Sea
of Azov (eastern population) (Pinchuk 1976; Pinchuk et al. 2003; Manilo & Peskov 2012). In waters of
Ukraine, the species also occurs in the southern part of the Dnipro Estuary, along the seashore from
Sychavka, Odesa Oblast to the mouths of the Berezansky and Dnipro–Bug Estuaries, in particular on
Dnister and Sanzheiska shoalbanks, in Odesa (Zambriborshch et al. 1995) and Karkinit Bay, as well
as near Cape Opuk (Shaganov 2009), in the Kerch Strait (Shaganov & Milovanov 2005), and in the
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southern part of the Sea of Azov to Cape Kazantip. Rarerly occurs in the northern part of the Sea of
Azov near Obitochna and Berdiansk Spits (Manilo 2009a, 2011b).
Ponticola eurycephalus (Kessler, 1874)
A species of the Ponto-Caspian faunal complex. Its geographic range covers the north-western part
of the Black Sea from the coast of Bulgaria along the coast of the Crimea to the Kerch Strait (Shaganov
& Milovanov 2005; Shaganov 2009; Eremeev et al. 2012), as well as the Sea of Azov from its southern
part to Obitochna and Berdiansk Spits, including estuaries of the Kuban region (Demchenko et al.
2005). In waters of Ukraine, the species was recorded in seaside waters of the Danube Delta (Balatskiy
& Voloshkevich 2005), in the Sasyk Estuary (Moshu 2006; Smirnov & Tkachenko 2007), and along the
coast of the Danube–Dnister interuve (Manilo 2009a). In the southern part of the Dnister Estuary,
it was identied as Neogobius cephalarges (Zambriborshch & Framudu 1990). e species also occurs
in coastal waters from the Dnister Estuary to the mouth of the Dnipro–Bug Estuary (Pinchuk 1977;
Shekk 2004); in Karkinit Bay it was mentioned as a species of the complex “G.cephalarges + platyro-
stris” (Pinchuk 1968; Manilo 2009a). According to some researchers, the north-western population
of the species is represented by the subspecies N. eurycephalus odessicus Pinchuk, 1977 (Manilo &
Peskov 2012).
Ponticola kessleri (Günther, 1861)
A species of the Ponto-Caspian faunal complex. Distributed in marine and freswaters of the basin
of the Black Sea (Svetovidov 1964; Kottelat & Freyhof 2007). e species is absent in the basin of the
Sea of Azov. In the last years, the species was recorded in the basin of the Rhine to which it entered
throught the Main–Danube Canal (Manné et al. 2013; Borcherding et al. 2011). In waters of Ukraine,
the species is known from lakes of the Danube Delta area such as Kytay, Katlabug, Yalpug, and Kagul
(Smirnov 1986). Also inhabits the Dnister to the mouth of its tributary Bystrytsia (Pinchuk 1977),
in the middle and lower courses of the Southern Bug and some of its tributaries (Movchan et al.
2003), as well as in the Dnipro to Kyiv (Smirnov 1986). e species was also recorded in the upper
section of the Sasyk (Moshu 2006) and Dnister Estuaries (Movchan et al. 2003), in the lower part of
the Khadzhibey Estuary (Kvach et al. 2009), and in the lower course of the Alma River in the Crimea
(Boltachev & Karpova 2017).
Ponticola platyrostris (Pallas, 1814)
A species of the Ponto-Caspian faunal complex. An endemic of the Black Sea known from the
western coast of the Crimea up to Georgia in the east; it was also found o the coast of Turkey
near Samsun (Vasileva 2007). Its records in checklists of shes of coastal waters of Bulgaria (Vassilev
& Pehlivanov 2005) and Romania (Radu & Radu 2008) belong to other goby species. In waters of
Ukraine, the species occurs near Cape Tarkhankut, in the west of the Crimea (Kacha) (Manilo 2009b),
as well as along the southern coast of the Crimea from Yalta to Karadag Nature Reserve (Vynogradov
1947) and Feodosia (Shaganov 2009). Rarerly occurs in the southern part of the Kerch Strait, prob-
ably only during exchange of saline waters from the Black Sea. e inclusion of the species into the
checklist of the ichthyofauna of the lower course of the Dnipro (Movchan 2001) and of the middle
course of the Southern Bug (Movchan et al. 2002) is erroneous.
Ponticola ratan (Nordmann, 1840)
A species of the Ponto-Caspian faunal complex. Its geographic range covers the basins of the
Black Sea (from the coast of Bulgaria to the western part of the Crimea and from the Kerch Strait to
Novorossiysk), the Sea of Azov (northern and eastern parts), and the Caspian Sea (Vasileva 2007).
In waters of Ukraine, the species was recorded near Zmiinyi Island (Snigiryov 2008), from the coast
of the Danube–Dnister interuve to Tendra Bay, including Shagan and Dnister shoalbanks (Manilo
2009a), in the southern part of the Dnister Estuary, in the Khadzhibey, Tiligul (from 1970 to 1990)
(Shekk 2004), Berezansky, and Dnirpo–Bug Estuaries, as well as in the Southern Bug to Bilousivka
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(Beling 1927). Pinchuk et al. (1985) mentioned an isolated dwarf population of the species in the
lower part of Kakhovka Reservoir near Berislav. In freshwaters, the species was found in Kamyanske
(Dniprodzerzhynsk) Reservoir nearby to the mouth of the Dnipro–Donbas Canal (Manilo &
Didenko 2013) and in Kaniv Reservoir (author’s unpublished data). e species was also recorded
near the Tarkhankut Peninsula (Pinchuk 1968; Eremeev et al. 2012), near Karadag and o the coast
of the Kerch Peninsula; it is abundant in the Kerch Strait (Shaganov 2009). e species is also known
from the southern part of the Sea of Azov to Cape Kazantip (Eremeev et al. 2012), in the Utlyuk
(Demchenko et al. 2005) and Molochnyi (during contact with the sea) Estuaries (Smirnov 2006), as
well as in the northern part of the sea to Berdyansk Spit.
Ponticola syrman (Nordmann, 1840)
A species of the Ponto-Caspian faunal complex. Its geographic range covers coastal waters of the
Black Sea (along the shore of Bulgaria, Romania, and Ukraine), the Sea of Azov, and the Caspian
Sea. Single records of the species are known from the Danube in Hungary (Guti 1999). In waters of
Ukraine, the species occurs in lakes of the Danube Delta area (Kytay, Katlabug, Yalpug and Kagul), in
seaside waters of the Danube Delta (Movchan et al. 2003), in the Dnister Estuary, and rarely along the
coast from the Dnister Estuary to the Grygorivsky Estuary (in some years during spring desalination)
(Vynogradov 1960; Zambriborshch et al. 1995). Earlier the species was recorded in the Tiligul (Shekk
2004), Berezansky, and Dnipro–Bug Estuaries (Pinchuk 1968; Movchan et al. 2003), and rarely in the
Kerch Strait (Shaganov & Milovanov 2005). In the Sea of Azov, the species is widely distributed and
also occurs in Lake Syvash (Demchenko 2005).
Genus Proterorhinus Smitt, 1899
Until recently, the genus was considered monotypic with a polymorphic species Proterorhinus
marmoratus (Pallas, 1814). Results of genetic studies (Slynko et al. 2012; Sorokin et al. 2011) indicate
that the Ponto-Caspian Basin is inhabited by two species of this genus: Pr.marmoratus represented
by populations in waters of the Black Sea, and Pr. semilunaris inhabiting saline and freshwaters of this
basin. In addition, the latter species also includes the earlier known species Pr. nasalis and Pr. semi-
pellucidus. e composition of the genus remains dubious and needs further clarication, although
currently it comprises 4–5 (?) species that inhabit fresh, brackish, and saline waters of the Azov–Black
Sea Basin, the Caspian Sea, as well as the north-eastern part of the Aegean Sea. In this review, we fol-
low the international database Fishbase (Froese & Pauly 2019) in which all these species are consid-
ered valid. us, in waters of Ukraine, four species of the genus occur.
Proterorhinus marmoratus (Pallas, 1814)
A species of the Ponto-Caspian faunal complex. Its geographic range is yet to be claried. In saline
and brackish waters of Ukraine, the species occurs in the coastal zone of the Black Sea near Zmiinyi
Island (Snigiryov 2008), in seaside waters of the Danube Delta (Balatskiy & Voloshkevich 2005),
along the coast of the Danube–Dnister interuve to the Grygorivsky Estuary, including the Budatsky,
Dnister, Tiligul (Shekk 2004), Berezansky, and Dnipro–Bug Eastuaries, as well as in Yahorlyk, Tendra,
and Dzharylhach Bays (Pinchuk 1987; Pinchuk & Tkachenko 1996; Movchan 2000; Manilo 2009a).
e species is known from nearby to Sevastopol and quite rarely also occurs along the southern coast
of the Crimea to the Kerch Strait (Bogutskaya & Naseka 2004; Shaganov & Milovanov 2005; Shaganov
2009). e species was recorded in the southern part of the Sea of Azov and in Lake Syvash, in the
Utlyuk and Molochny Estuaries, as well as near Berdiansk Spit (Demchenko & Mityai 2001; Diripasko
et al. 2011; Demchenko et al. 2005).
Proterorhinus nasalis (De Filippi, 1863)
A species of the Ponto-Caspian faunal complex. Its geographic range covers the basins of the Sea of
Azov and the Caspian Sea. In Ukraine, the species occurs in the coastal desalinated zone of the Sea of
Azov and is known from the Molochny Estuary, near Obitochna and Berdiansk Spits, as well as from
Lake Syvash (Movchan 2011). e species’ range needs to be claried.
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Proterorhinus semilunaris (Heckel, 1837)
A species of the Ponto-Caspian faunal complex. It is distributed in estuaries of the western and
north-western parts of the Black Sea and in rivers that ow into them. e species is known in the
Danube from its mouth to Central and Western Europe (Manne & Poulet 2008; Manne et al. 2013), in
the basin of the Dnister, in the Southern Bug and Dnipro Rivers, in the lower course of the Marytsia
and Struma Rivers that ow into the eastern part of the Aegean Sea (Kottelat & Freyhof 2007; Froese
& Pauly 2019), as well as in the basin of the Vistula to its mouth (Grabowska et al. 2008). e species
was brought to lakes and rivers of North America with ballast waters (Stepien & Tumeo 2006; Neilson
& Stepien 2009).
Proterorhinus tataricus Freyhof et Naseka, 2007
A species of the Ponto-Caspian faunal complex. A strict endemic of freshwaters of the Crimea. It
is known from the Chorna River (Freyhof & Naseka 2007), which ows into Sevastopol Cove.
Subfamily Gobiinae
Genus Aphia Risso, 1827
A monotypic genus.
Aphia minuta (Risso, 1810)
A species of the Atlantic-Mediterranean faunal complex. Distributed in the eastern Atlantic from
the Scandinavian Peninsula to Morocco. Also inhabits the Mediterranean (not recorded along the
northern coast of Africa), Adriatic, Aegean, Marmara, and Black Seas, and occurs in the southern part
of the Sea of Azov (Svetovidov 1964; Vasileva 2007). In waters of Ukraine, the species was recorded in
seaside waters of the Danube Delta (Chepurnov et al. 1954), in the Sasyk Estuary before its separation
from the sea by a dam and desalination by waters from the Danube (Smirnov & Tkachenko 2007),
along the seashore in the north-western part of the Black Sea to Odesa (Kvach 2015), in Karkinit Bay
in the area of Zernov’s Phyllophora Field Botanical Sanctuary, in Yahorlyk and Tendra Bays, near the
Tarkhankut Peninsula (Vynogradov 1960), near Zmiinyi Island (Snigiryov 2008), in coastal waters of
north-western Crimea (Chesalin et al. 2004; Karpova 2009) and Karadag (Vinogradov 1949), in the
Kerch Strait (Shaganov & Milovanov 2005), and in the southern part of the Sea of Azov (Diripasko et
al. 2011). One of the few pelagic species of the family.
Genus Chromogobius de Buen, 1930
e genus consists of three species distributed in the eastern Atlantic and in the Mediterranean
and Black Seas. Two species occur in waters of Ukraine.
Chromogobius quadrivittatus (Steindachner, 1863)
A species of the Atlantic-Mediterranean faunal complex. Its geographic range covers coastal wa-
ters of the northern (Spain, France, Italy, Croatia) and eastern parts of the Mediterranean Sea (Adib
2005). Also occurs in the Sea of Marmara, the Aegean (Bilecenoglu et al. 2014) and the Black Seas.
A highly rare species in every part of the range. In the Black Sea, it was earlier also known near the
coast of the Caucasus (Pinchuk 1987), where by know has entirely disappeared, as well as in Varna
Bay (Georgiev 1961). A single record of the species during sh kills near Odesa is reported by A. K.
Vinogradov and S. A. Khutornoy (2013), although the specimen was not preserved and only its pho-
tograph was published. In waters of Ukraine, the species was rst recorded in underwater karst caves
of Cape Tarkhankut (Kovtun 2013). In 2015, a number of specimens, including juveniles, were found
in Martynova Cove (Sevastopol, Crimea) (Boltachev & Karpova 2016), and in Kozacha Cove in 2016
(Boltachev & Karpova 2017). According to the latest data, the species was also revealed in underwater
caves of Bulgaria north of Varna Bay nearby to the border with Romania.
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Chromogobius zebratus (Kolombatović, 1891)
A species of the Atlantic-Mediterranean faunal complex. A recent invader of the Mediterranean. It
is known from several localities of the northern (Spain, France, Italy) (Colombo & Langeneck 2013)
and eastern parts of the Mediterranean (Lebanon, Turkey) (Engin & Dalgiç 2008), Adriatic (Croatia,
Montenegro), and Aegean Sea (Greece) (Miller 1986). A rare and non-abundant species. In the Black
Sea, it was rst recorded in waters of Ukraine in underwater caves of Cape Tarkhankut (Kovtun &
Karpova 2014; Boltachev & Karpova, 2014, 2017).
Genus Gammogobius Bath, 1971
A monotypic genus.
Gammogobius steinitzi Bath, 1971
A species of the Atlantic-Mediterranean faunal complex. Known from several localities of the
northern part of the Mediterranean Sea: o the coast of France (Dufour et al. 2007), near Ibiza,
(Scsepka & Ahnelt 1999), in the north of the Tyrrhenian Sea near Giglio Island, Italy (Ahnelt et al.
1998), in the north of the Adriatic Sea near Krk, Croatia (Kovačić 1999), as well as near Crete, Greece
(Kovačić et al. 2011). A rare and non-abundant species. It is known in the Black Sea only in waters
of Ukraine from marine underwater karst caves of the Tarkhankut Peninsula near Malyi Atlesh tract
(Kovtun 2012; Kovtun & Manilo 2013).
Genus Gobius Linnaeus, 1758
e genus includes 28 species distributed in the Indo-West Pacic, Atlantic Ocean, Mediterranean
Sea, Black Sea, and the Sea of Azov. In waters of Ukraine, currently seven species are known.
Gobius bucchichi Steindachner, 1870
A species of the Atlantic-Mediterranean faunal complex. Distributed in the eastern Atlantic from
the south of Portugal to the coasts of Morocco. Also occurs in the Mediterranean Basin: in the Adriatic,
Aegean, and Marmara Seas. In the Black Sea, the species is distributed along the coasts of Turkey, the
Caucasus, the Crimea, and Bulgaria (Vasileva 2007). In Ukraine, the species inhabits coastal waters
along the western shore of the Crimea near Cape Tarkhankut and Sevastopol (Boltachev & Karpova
2012b; Manilo 2014).
Gobius cobitis Pallas, 1814
A species of the Atlantic-Mediterranean faunal complex. Distributed in the eastern Atlantic
from the English Channel to Morocco, as well as in the Mediterranean Basin (Adriatic, Aegean, and
Marmara Seas). In the Black Sea, the species occurs near the coasts of the Crimea, the Caucasus,
Turkey, Romania, and Bulgaria. Absent in the Sea of Azov (Vasileva 2007). In waters of Ukraine, it
was recorded near Cape Tarkhankut (Malyi Atlesh tract), along the southern coast of the Crimea
from Sevastopol to Karadag and Feodosia (Boltachev 2003; Gordina et al. 2004; Shaganov 2009). In
the north-western part of the Black Sea, it was discovered near Zmiinyi Island (Manilo 2009a).
Gobius couchi Miller et El-Tawil, 1974
A quite rare Atlantic-Mediterranean species. A recent invader of the Mediterranean Basin. It
was described from coastal waters of Great Britain (Miller & El-Tawi l 1974) and later recorded near
south-western and northern Ireland (Minchin 1988; Costello 1992). Later, the species was found in
the Mediterranean Basin: in 1996–1997, near the coast of Croatia (Kovačić 2001); in 1999, in the
Tyrrhenian Sea near Ischia (Stefanni & Mazzoldi 1999); in 2004, in the Ionian Sea near Corfu (Šanda
& Kovačić 2009); in 2007, in the Aegean Sea, in the southern part of the Dardanelles (Ozen et al.
2009); in 2009, in the Ligurian Sea (Liu et al. 2009); in 2010, near Crete (Kovačić et al. 2011); in 2011,
near Malta (Kovačić et al. 2013). In the Black Sea, the species was recorded in 2015–2016 only in
waters of Ukraine in Kozacha and Karantynna Coves (Karpova & Boltachev 2018), where the species
has completely adapted to conditions of the south-western coast of the Crimea.
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Gobius cruentatus Gmelin, 1789
A species of the Atlantic-Mediterranean faunal complex. A recent invader to the Mediterranean.
Widely distributed in the eastern Atlantic from south-western Ireland to Morocco and Senegal, as
well as in seas of the Mediterranean Basin (Adriatic, Aegean, Marmara Seas, and the Bosporus) and
in the Black Sea (Golani et al. 2006). Its rst records in the Black Sea are conned to coastal waters
of Ukraine near Sevastopol in Martynova Cove from 2002. Later, in 2007, a number of specimens
were caught o the coast and in coves of Sevastopol (Boltachev et al. 2009a; Boltachev & Karpova
2014). About the same time, the red-mouthed goby was recorded near the coast of Turkey (Engin et
al. 2007). Since then a stable tendency of increase in abundance and distribution has been observed
along the south-western coast of the Crimea to Laspi Cove (Boltachev & Karpova 2014).
Gobius niger Linnaeus, 1758
A species of the Atlantic-Mediterranean faunal complex. Widely distributed in waters of the east-
ern Atlantic from the coast of Norway and the Baltic Sea to Morocco, including the Canary Islands.
A common species in the Mediterranean, Adriatic, Aegean, and Marmara Seas. In the Black Sea, the
species occurs along the entire coast (Vasileva 2007). S. P. Volovik and S.A.Chikhachev (1998) noted
the expansion of the species’ range in the Sea of Azov to its northern parts. In waters of Ukraine, the
species occurs near Zmiinyi Island (Snigiryov 2008), in seaside waters of the Danube Delta (also oc-
curred in the Sasyk Estuary before its desalination (Smirnov & Tkachenko 2007)), along the coast
from the Danube–Dnister interuve to the Grygorivsky Eastuary including Shagan and Dnister
shoalbanks, in the Budak Estuary, in the southern part of the Dnister Estuary (Manilo 2009a), in
the Tiligul Estuary (Shekk 2004), at the mouth of the Dnipro–Bug Estuary, as well as in Tendra,
Yahorlyk, Dhzarylhach, and Karkinit Bays (Pinchuk & Tkachenko 1996) and near the coast of the
Crimea (Gordina et al. 2004; Shaganov 2009).
Gobius paganellus Linnaeus, 1758
A species of the Atlantic-Mediterranean faunal complex. Distributed in the eastern Atlantic from
the western coast of Scotland to Senegal, including the Azors, the Canary Islands, and Madeira. Also
occurs in the Mediterranean, Adriatic, Aegean, and Marmara Seas. rought the Suez Canal, it pen-
etrated the Red Sea. In the Black Sea, the species is known from coastal waters of Bulgaria, Turkey,
the Caucasus, and the south of the Crimea (Vasileva 2007). In waters of Ukraine, the species occurs
near Zmiinyi Island (Snigirov et al. 2018), Sevastopol, and the area of the Tarkhankut Peninsula, as
well as along the southern coast of the Crimea to Feodosia where it is a rare species (Boltachev 2003;
Shaganov 2009).
Gobius xanthocephalus Heimer et Zander, 1992
A species of the Atlantic-Mediterranean faunal complex. A recent invader of the Mediterranean
Sea. Inhabits coastal waters of the eastern Atlantic along the shores of Portugal (Almeida & Arruda
1998) and the Canary Islands (Wirtz & Herrera 1995). In the Mediterranean Sea, it is known o the
coast of France (Heimer & Zander 1992). e species’ occurrence in the central and eastern parts of
the Mediterranean Sea (from Italy to Turkey) is debatable (Louisy 2002). It is a rare species in the
Black Sea, where it was recorded in its eastern part near Abkhazia (Vasileva & Bogorodskii 2004). In
waters of Ukraine, the species was found in the Crimea in coves of Sevastopol (Boltachev et al. 2009а)
and near Cape Tarkhankut (Karpova & Saksahanskyy 2011).
Genus Knipowitschia Iljin, 1927
e genus includes 17 species distributed in basins of the Mediterranean, Black, and Caspian Seas.
Two species are known in saline, brackish, and freshwaters of Ukraine.
Knipowitschia caucasica (Berg, 1916)
A species of the Atlantic-Mediterranean faunal complex. e most widely distributed representa-
tive of the genus occurring in basins of the Sea of Azov, the Black, Caspian, Ionian, Adriatic, Aegean,
and Marmara Seas (Economidis & Miller 1990; Ahnelt et al. 1995). In the 1950s, it was accidently
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introduced in the Aral Sea, where by now its population diasappeared. In the Black Sea, the species
occurs in the western and northern parts along the coasts of Bulgaria (in lakes near Burgas and in the
lower course of rivers that ow into them) (Vassilev & Pehlivanov 2005), Romania (Banarescu 1964),
and further to the Kerch Peninsula and the coast of the Caucasus. In marine and brackish waters of
Ukraine, the species was recorded in the Danube Delta (Balatskiy & Voloshkevich 2005), in the Sasyk
Estuary before its separation from the sea by a dam (Smirnov & Tkachenko 2007), along the shore
of the Danube–Dnister interuve (Chepurnov 1958), in the Budatsky (Manilo 2014), Tiligul (Shekk
2004), and Berezansky Estuaries, in Tendra Bay (Vinogradov et al. 1967; Manilo 2009a), in the Kerch
Strait, in the Eastern Syvash, in the Utlyuk and Molochnyi Estuaries (Demchenko et al. 2005), and in
mouths of small rivers (Diripasko et al. 2011). According to A. V. Pankov (Pankov 2007), the species is
actively expanding its range to the north in the Dnipro River. It was also recorded in the upper course
of the Siversky Donets (tributary of the Don) within Kharkiv Oblast, Ukraine (Shandikov et al. 2009).
Knipowitschia longecaudata (Kessler, 1877)
A species of the Atlantic-Mediterranean faunal complex. Distributed sporadically in the western
and south-western parts of the Black Sea (Bulgaria, Romania, and Ukraine), in the southern and
eastern parts of the Sea of Azov, and in the northern part of the Caspian Sea (Troitskiy & Tsunikova
1988). It was also recorded in Turkey in Lake Manyas (Turan et al. 2005). In waters of Ukraine, the
species was discovered in lakes of the Danube Delta area (Zelenin & Vladimirov 1975), in the lower
part of the Danube Delta (Balatskiy & Voloshkevich 2005), in the Sasyk (Moshu 2006; Smirnov &
Tkachenko 2007), Dnister, Tiligul (Shekk 2004), and Dnipro–Bug Estuaries and in the lower course
of rivers that ow into them. In P. J. Miller’s opinion (Miller 2004), records of the species in the
Crimea near Sevastopol and Yalta and in the Yahorlyk Esturary (Vinogradov 1960) possibly belong to
Pomatoschistus marmoratus. e only specimen was caught in the estuary zone of the Chorna River
near Sevastopol (Boltachev & Karpova 2012a). In the area of the Kerch Strait, records of the species
were reported from desalinated estuaries of the Kuban region. It was also found in the North Crimean
Canal (Karpova 2009). A semipelagic species.
Genus Millerigobius Bath, 1973
A monotypic genus.
Millerigobius macrocephalus (Kolombatovič, 1891)
A species of the Atlantic-Mediterranean faunal complex. A recent invader of the Mediterranean
Basin. It was known as an endemic of the Aegean and Adriatic Seas (Miller 1986; Golani et al. 2006;
Kovačić 2008). It was also found near Ibiza (Fischer et al. 2007), Corsica, and near the entrance to the
Dardanelles (Bogorodsky et al. 2010). A rare species. It was not recorded in the Black Sea earlier. In
waters of Ukraine, the species was rst discovered in April 2009 in Sevastopol Cove during inspection
of articial mussel collectors installed at the mouth of the cove (Boltachev et al. 2010). By now, the
species has formed here a separate population.
Genus Pomatoschistus Gill, 1863
e genus includes 14 species that inhabit the coastal zone from the north-eastern Atlantic to the
Black Sea and Sea of Azov. ree species are known in marine and brackish waters of Ukraine.
Pomatoschistus bathi Miller, 1982
A species of the Atlantic-Mediterranean faunal complex. e native range of the species covers the
Mediterranean, Aegean, and Marmara Seas. In the Black Sea, it is known near the coast of Abkhazia
(Vasileva & Bogorodskii 2004), and near the Black Sea coast of Russia (Vasileva 2007) and Bulgaria
(Vassilev et al. 2012). In marine waters of Ukraine, the species was rst recorded in the early 2000s
in coves of Sevastopol and along the southern coast of the Crimea from Cape Sarych to Cape Tovsty
(Boltachev & Karpova 2010). Currently, the species is abundant in coves of Sevastopol and Lake
Donuzlav, as well as near the Tarkhankut Peninsula (Eremeev et al. 2012). Also occurs along the
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southern coast of the Crimea (it was recorded in 2013 in the Karadag Nature Reserve) (Boltachev et
al. 2016) and in Karkinit Bay (Prishchepa et al. 2018). In A. R. Boltachev and E. P. Karpova’s (2010)
opinion, a self-maintaining population of the species has formed in the coastal zone of the Crimea.
Pomatoschistus marmoratus (Risso, 1810)
A species of the Atlantic-Mediterranean faunal complex. Inhabits coastal waters of the eastern
Atlantic from Great Britain to the Bay of Biscay, Portugal, and Spain (Froese & Pauly 2019). Also oc-
curs in the Mediterranean, Aegean, and Marmara Seas, and in the Suez Canal. It was introduced to
Lake Qarun, Egypt (Vasileva 2007). In the Black Sea and Sea of Azov, it is widely distributed along al-
most the entire coast. In waters of Ukraine, it occurs along the entire coast in marine waters, estuaries,
saline and brackish lakes (Gordina et al. 2004; Shekk 2004; Demchenko 2005; Shaganov & Milovanov
2005; Manilo 2009a; Movchan 2011).
Pomatoschistus minutus (Pallas, 1770)
A species of the Atlantic-Mediterranean faunal complex. Inhabits coastal waters of the eastern
Atlantic from the coast of Norway to the southern part of the Iberian Peninsula; also common in the
Mediterranean, Aegean, Marmara, and Black Seas (Miller 1986; Vasileva 2007). In waters of Ukraine,
it was recorded in coastal waters of Zmiinyi Island (Pinchuk & Savchuk 1982), in seaside waters of the
Danube Delta, along the coast of the Danube–Dnister interuve, in the Budatsky Estuary, in the lower
part of the Dnister Estuary, in Odesa Bay (Kvach 2015), at the mouth of the Berezansky Estuary, in
Yahorlyk, Tendra, and Dzharylhach Bays (Vynogradov 1960; Pinchuk & Tkachenko 1996; Movchan
2000), as well as in the Crimea in Lake Donuzlav (Boltachev & Zuyev 1999), near Sevastopol (Karpova
2009), and in the Kerch Strait. e species was not noted for the Sea of Azov until the mid-1950s,
although in the mid-1970s it has expanded its range to the northern part of sea (Pinchuk & Savchuk
1982). Eventually, the species was included into the checklist of shes of the Sea of Azov (Diripasko
et al. 2011).
Genus Zebrus de Buen, 1930
A monotypic genus.
Zebrus zebrus (Risso, 1827)
A species of the Atlantic-Mediterranean faunal complex. Distributed in the Mediterranean Sea ex-
cept for small areas of the northern coast of Africa (Kovačić et al. 2005; Bogorodsky et al. 2010). In the
eastern Atlantic, it was recorded near the coast of Spain in the southern part of the Iberian Peninsula
(Froese & Pauli 2019). A recent invader of the Mediterranean Sea. In 2007, a mature specimen was
caught in the Black Sea o the coast of Turkey near Cape Jason (Kovačić & Engín 2009). In waters of
Ukraine, the species was rst recorded near the shore of the Crimea at mussel collectors in Sevastopol
Cove, where since 2009 co-occurs with M. macrocephalus allowing to suggest that the species has
naturalised in the Crimea (Manilo et al. 2013а; Karpova et al. 2015).
Genus Zosterisessor Whitley, 1935
A monotypic genus.
Zosterisessor ophiocephalus (Pallas, 1814)
A species of the Atlantic-Mediterranean faunal complex. Distributed along the northern and
south-western coast of the Mediterranean Sea, as well as in the Aegean and Marmara Seas (Miller
1986). Its records in the Bay of Biscay are dubious (Pinchuk et al. 2004b). In the Black Sea, the species
is known along almost the entire shore except for the central part of the northern coast of Turkey,
and it is very rare near the southern coast of the Crimea. In the Sea of Azov, it is distributed from
the south-western to the northern coast (Demchenko et al. 2005). In waters of Ukraine, the species
occurs from seaside waters of the Danube Delta to the Kerch Strait, including all estuaries and bays.
In the Sasyk Estuary, the species disappeared aer its separation from the sea by a dam (Smirnov &
Tkachenko 2007).
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Discussion
In summary, we can note that the true gobies are currently represented in Ukraine by 38species of
17genera. e fauna of the family Gobiidae includes representatives of the Ponto-Caspian (20 spe-
cies, 52.6 % of the total number of species of the family in Ukraine) and Atlantic-Mediterranean
(18 species, 47.4 %) faunal complexes (see Table 1).
Species Faunal complex
Benthophiloides brauneri PC
Benthophilus magistri PC
B. nudus PC
B. stellatus PC
Caspiosoma caspium PC
Neogobius uviatilis PC
N. melanostomus PC
Babka gymnotrachelus PC
Mesogobius batrachocephalus PC
Ponticola cephalargoides PC
P. eurycephalus PC
P. kessleri PC
P. platyrostris PC
P. r at an PC
P. s yr ma n PC
Proterorhinus marmoratus PC
P. nasalis PC
P. semilunaris PC
P. tataricus PC
Aphia minuta AM
Table 1. The current species composition of the family Gobiidae of waters of Ukraine and the faunal com-
plexes to which they belong
Таблиця 1. Сучасний видовий склад родини Gobiidae водойм України та їх належність до фауністич-
них комплексів
Species Faunal complex
Chromogobius quadrivittatus* AM
C. zebratus*? AM
Gammogobius steinitzi*? AM
Gobius bucchichi AM
G. cobitis AM
G. couchi* AM
G. cruentatus* AM
G. niger AM
G. paganellus AM
G.
xanthocephalus*
AM
Knipowitschia caucasica АМ
K. longecaudata PC
Millerigobius macrocephalus* AM
Pomatoschistus bathi* AM
P. marmoratus AM
P. minutus AM
Zebrus zebrus* AM
Zosterisessor ophiocephalus AM
Total (n) 20 18
(%) 52.6 47.4
Note. Faunal complexes: PC — Ponto-Caspian, AM — Atlantic-Mediterranean, * — recent immigrant species; *?— time
of appearance unknown; endemic species are given in bold.
Among species of the Atlantic-Mediterranean complex, a group of recent invaders can be high-
lithed, which appeared in the Black Sea in the early 2000s (Chromogobius zebratus, Gobius couchi,
G. cruentatus, G. xanthocephalus, Millerigobius macrocephalus, Pomatoschistus bathi, and Zebrus ze-
brus). is is directly related to the natural permanent process of Mediterranization. e time of ap-
pearance of two species (Chromogobius zebratus and Gammogobius steinitzi) remains unknown. By
now, near the coast of Ukraine and in coves of the Crimea such species as C. quadrivittatus (accord-
ing to records of juveniles in coves of Sevastopol), G. cruentatus, M. macrocephalus, and P. b ath i have
fully naturalised. Four species of the family Gobiidae are endemic to Ukraine: Benthophilus magistri,
B. stellatus, P. platyrostris, and Proterorhinus tataricus.
References
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