Alien fishes in the Black Sea waters of Crimea (Ukraine)

Vector of introduction. A bentho-pelagic species that have spread to the coast of Crimea indi-vidually through the Bosphorus from the Marmara Sea (mediterranisation).

Order Pleuronectiformes Family Scophthalmidae Jordan, 1923 Genus Scophthalmus Rafinesque, 1810

24. Scophthalmus rhombus (Linnaeus, 1758) -- Brill

Synonyms: Pleuronectes rhombus Linnaeus, 1758; Pleuronectes laevis Turton, 1802; Rhombus laevis (Turton, 1802).

Material (in the fish collection of NMNH NASU): absent.

Brief description. D 83, A 61, P 11-12, V 6, C 17, sp.br. 18 (Giragosov et al. 2012). Distribution. The species is distributed in the eastern part of the Atlantic Ocean from Norway and Great Britain to the north of Africa, also occurs in the Baltic, Mediterranean, Aegean, and Mar-mara Seas and in the Bosphorus (Mengi 1971; Nielsen 1986). In the northern part of the Black Sea, only 3 specimens have been recorded for the entire history of ichthyological research: 1 near Feodosia in the 19th century (Kessler 1877) and 2 near Kara-Dag in 1946 (Vinogradov 1949). A female specimen was found off the shore of Bulgaria in May 1954 (Georgiev et al. 1960). In 2010, 3 specimens were caught by seine in Striletska Cove in Sevastopol (Giragosov et al. 2012).

Vector of introduction. A typical benthic species with no long-term migrations, although it clearly apparead near Crimea naturally through the Bosphorus from the Marmara Sea (mediterranisation).

Species that occur casually (single records) in waters of Crimea

Order Gadiformes Family Gadidae Rafinesque, 1815 Genus Micromesistius Gill, 1863

25. Micromesistius poutassou (Risso, 1827) -- Blue whiting

Synonyms: Merlangus poutassou Risso, 1827; Boreogadus poutassou (Risso, 1827); Gadus poutassou (Risso, 1827); Merlangus vernalis Risso, 1827; Merlangus pertusus Cocco, 1829.

Material (in the fish collection of NMNH NASU): absent.

Brief description. D1 13 (12-14), D2 14 (12-14), D3 23 (23-26), A1 35 (33-39), A2 26 (24-27), P 22, sp. br. 27-33 (Movchan 2011).

Distribution. An Atlantic-Boreal species distributed from Spitsbergen along the coast of Europe to the Canary Islands, northwest Africa to Cape Bojador, as well as to west from the Barents Sea to Iceland, Greenland, and northeast coast of the USA. The species inhabits most seas of the Mediterranean Basin, although it is more common in the western part of the Mediterranean Sea (Svetovidov 1964; Marine Species... 2000). The only specimen known in the Black Sea was caught by hook in January 1999 near Cape Aya (southwest coast of Crimea) at a depth of 60 m (Boltachev et al. 1999, 2010).

Vector of introduction. Accidental natural spread from seas of the Mediterranean Basin through the Bosphorus (mediterranisation).

Order Scorpaeniformes Family Dactylopteridae Gill, 1885 Genus Dactylopterus Lacepede, 1801

26. Dactylopterus volitans (Linnaeus, 1758) -- Mediterranean flying gurnard

Synonyms: Trigla volitans Linnaeus, 1758; Dactylopterus vulgaris Steindachner, 1867; Cephalacanthus volitans (Linnaeus, 1758); Dactylopterus blochii Swainson, 1839.

Material (in the fish collection of NMNH NASU): No. 4970.

Brief description. D1 VII; D2 VIII; A 6 (our data).

Distribution. The species is distributed in the eastern and western Atlantic Ocean south of the English Channel to Angola and from Massachusetts (USA) to Argentina, as well as near Madeira, the Azores, and in seas of the Mediterranean Basin (Mediterranee... 1987; Froese & Pauly 2000). In the Black Sea, 2 records of the species were reported from Odessa Bay in September 1979. (Movchan 2011) and an immature 40 mm long specimen from near Sevastopol (Liubymivka village) (Boltachev & Karpova 2014).

Vector of introduction. It is hard to determine the species' vector of appearance; it could be passive spread by either currents or ballast waters.

Order Perciformes

Family Sparidae Bonaparte, 1832

Genus Dentex Cuvier, 1814

27. Dentex dentex (Linnaeus, 1758) -- Common dentex

Synonyms: Sparus dentex Linnaeus, 1758; Dentex vulgaris Valenciennes, 1830.

Material (in the fish collection of NMNH NASU): absent.

Brief description. D X-XII 11; A III 8; P I 14; V I 5; l.l. 60-65 (Movchan 2011).

Distribution. The species is distributed in the eastern Atlantic Ocean from the Bay of Biscay and southern part of the British Isles to Senegal, near Madeira and the Canary Islands, as well as in the Mediterranean, Aegean, and Marmara Seas. Rare records of the species are known from the Black Sea off the coast of Turkey, Bulgaria, and Romania. In waters of Ukraine, only a single specimen was caught in June 2014 in the coastal zone of Cape Fiolent at a depth of 9 m (Boltachev & Karpova 2017).

Vector of introduction. Independent accicental spread from seas of the Mediterranean Basin through the Bosphorus (mediterranisation).

Family Chaetodontidae Rafinesque, 1815

Genus Heniochus Cuvier, 1816

28. Heniochus acuminatus Linnaeus, 1758 -- Pennant coralfish

Synonyms: Chaetodon acuminatus Linnaeus, 1758; Chaetodon macrolepidotus Linnaeus, 1758; Heniochus macrolepidotus

(Linnaeus, 1758); Taurichthys macrolepidotus.

Material (in the fish collection of NMNH NASU): No. 8099, 10695.

Brief description. D XI-XII 22-27, A III 17-19, P I 16, V I 5, l.l. 51 (our data).

Distribution. Tropical waters of the Indian and Pacific Oceans from South Africa to Japan and Papua New Guinea. The only finding of the species in the Black Sea was reported in October 2003 from Balaklava Bay (Boltachev & Astakhov 2004).

Vector of introduction. An Indo-West Pacific species, possibly spread by ballast waters.

Family Acanthuridae Bonaparte, 1835

Genus Acanthurus Forsskal, 1775

29. Acanthurus monroviae Steindachner, 1876 -- Monrovia doctorfish

Synonyms: Teuthis munroviae Steindachner, 1876; Acanthurus monrovie Steindachner, 1876; Acanthurus phlebotomus

(non Valenciennes, 1835).

Material (in the fish collection of NMNH NASU): absent.

Brief description. D IX 24-26, A VI 24-26 (Froese & Pauly 2020).

Distribution. The native range of the species covers the eastern Atlantic Ocean from the coast of Morocco to Angola, the Canary Isalnds, and the islands of Cap-Vert. A significant increase in the number of records of A. monroviae and in the expansion of its range has been observed since the 2000s in the Mediterranean Sea: specimens have been found off the coast of southern Spain, Malta, Tunisia, Algeria, Israel, and Greece. The only specimen with TL 235 mm known from waters of Ukraine was caught on 19 September 2015 by fishing net near the entrance of Balaklava Bay in the southwest part of the Crimean Peninsula (Boltachev & Karpova 2020).

Vector of introduction. A bentho-pelagic species that appeared in coastal waters of Crimea from eastern seas of the Mediterranean Basin through the Bosphorus (mediterranisation).

Family Sphyraenidae Rafinesque, 1815

Genus Sphyraena Rцse, 1793

30. Sphyraena pinguis Gьnther, 1874 -- Red barracuda

Synonyms: absent.

Material (in the fish collection of NMNH NASU): No. 6525.

Brief description. D1 V, D21 9, A II 9, P 11-13, VI 5, l.l. 80-92 (our data).

Distribution. A pelagic species migrating in flocks. It is widely distributed in the Indo-West Pacific region along the continental shelf of East Africa from the Red Sea to Mozambique and the Republic of South Africa, as well as along the shores of Asia from the Arabian Peninsula to Indonesia. The species also occurs in the Pacific Ocean from the southern Primorye and Hokkaido to Papua New Guinea and southern Australia (Doiuchi & Nakabo 2005). Through the Suez Canal, the species spread to the eastern part of the Mediterranean Sea (Lessepsian migrant) and it was recorded off the coast of Palestina in 1931 (Golani et al. 2002). In following years, the red barracuda has spread along the coast of Egypt, Israel, Lebanon, and Turkey, including the Aegean Sea. The species also occurs near Malta and it has completely naturalised in the eastern part of the Mediterranean Sea (Golani et al. 2002). Two specimens were caught in Balaklava Bay in August 1999, which were initially identified as S. obtusata Cuvier, 1829, but later re-identified as S. pinguis (Boltachev 2009).

Vector of introduction. The species' vector of introduction near the coast of Crimea is migra-tion from the Marmara Sea through the Bosphorus (mediterranisation).

Order Tetraodontiformes

Family Tetraodontidae Rafinesque, 1815

Genus Lagocephalus Swainson, 1839

31. Lagocephalus sceleratus Gmelin, 1789 -- Silver-cheeked toadfish

Synonyms: Fugu sceleratus (Gmelin, 1789); Gastrophysus sceleratus (Gmelin, 1789); Tetraodon sceleratus Gmelin, 1789;

Tetraodon bicolor Brevoort, 1856; Tetraodon blochii Castelnau, 1861.

Material (in the fish collection of NMNH NASU): No. 4005, 4961, 3117, 4196.

Brief description. Only soft radials in the fins, 10-13 in the dorsal and 8-12 in the anal fin (our data).

Distribution. The species is distributed in coastal waters of the tropical zone of the Indian and Pacific Oceans. It has spread to the Mediterranean Basin through the Suez Canal (Lessepsian migrant), where it was first recorded in 2003 in the Aegean Sea. Currently, the species has widely dispersed in the eastern and central parts of the Mediterranean Sea near the coast of Israel, Turkey, and Greece, as well as in the Aegean Sea to the Dardanelles. A living specimen was caught by net in Staropivnichna Cove (Sevastopol Bay) in November 2014 (Boltachev & Karpova 2017).

Vector of introduction. The species' vector of introduction near the coast of Crimea is migra-tion from the Marmara Sea through the Bosphorus (mediterranisation).

Among the introduced species that occur regularly in waters of Crimea, benthic species pre-dominate (12 species, or 80.0%), while bentho-pelagic and pelagic species make up an insignificant part--6.7% and 13.3%, respectively. Among species that occur sporadically, pelagic (50.0%) and bentho-pelagic species prevail (37.5%). Bentho-pelagic species predominate among fishes that occur casually (57.1%) (Fig. 2).

Fig. 2. The ratio of species of different ecological groups (benthic, bentho-pelagic, pelagic) among alien fishes in the Black Sea waters of Crimea.

Рис. 2. Співвідношення видів різних екологічних груп (бентичної, бентопелагічної, пелагічної) серед риб-вселенців в чорноморських водах Криму.

Among the 31 introduced species recorded in the Black Sea waters of Crimea for the past 50 years, species that appeared due to the natural process of mediterranisation prevail (22 species, or 71.0%). The vector of introduction could not be identified for 4 species (Syngnathus acus, Dactylopterus voli- tans, Sparus aurata, and Parablennius incognitus), the ratio of which is 13.3%. The appeareance of the remaining 5 species (Planiliza haematocheila, Sebastes schlegelii, Heniochus acuminatus, Millerigobius macrocephalus, and Tridentiger trigonocephalus), which make up 16.1%, is due to anthropogenic factors (Fig. 3).

Fig. 3. The ratio of alien fishes in the Black Sea waters of Crimea with different vectors of introduction.

Рис. 3. Співвідношення видів-вселенців в чорноморських водах Криму з різними векторами вселення.

Note. For vectors of introduction, see Table. 1 (grey -- introduction related to mediterrani- sation, white -- unknown vector of introduction, black -- introduction due to anthropogenic factors).

Примітка. Позначення векторів вселення див. у табл. 1 (сірий -- поява пов'язана з процесом медітерранізації, білий -- невідомий вектор вселення, чорний -- поява через антропогенні фактори).

In general, benthic species predominate among the introduced fishes (14 species, or 45.2%), whereas bentho-pelagic (9) and pelagic (8) species make up only 29.0% and 25.8%, respectively.

Of the 31 introduced species in marine waters of Crimea, 10 have naturalised and acclimatised (32.2%), forming independent breeding population. Among them, species of the family Gobiidae (Ch. quadrivittatus, G. couchi, G. cruentatus, G. xanthocephalus, M. xanthocephalus, P. bathi, and Z. zebrus) and the taxonomically close family Oxudercidae (T. trigonocephalus) significantly prevail, whereas one species P incognitus represents the family Blenniidae. The only representative of the pelagic ecological group of fishes from the family Mugilidae -- the redlip mullet P haematocheila -- was intentionally introduced in the 1970s and it has completely acclimatised in the Azov-Black Sea Basin, including coastal waters of Crimea, and has become an important commercial object.

Conclusions

The main vector of introduction of new species of fish in the Black Sea is natural migration, as well as the spread of juveniles through the Bosphorus by currents (mediterranisation) and the Black Sea plays the role of recipient area. Further dispersal of species to coastal waters of Crimea is related to the geographic location of the peninsula, currents, habitat diversity, and biological features of species.

The appearance of alien fish species in coastal waters of Crimea is currently considered as an increasing invasion process. For the past 50 years, 31 species have been recorded in this region, 48.4% of which occur regularly, 29.0% sporadically, and 22.6% casually. Of the 10 species that have naturalised and acclimatised, only one was introuduced intentionally.

Benthic and bentho-pelagic species predominate among the introduced fishes.

The increase of species diversity of fishes in the Azov-Black Sea region (including Crimea) due to alien species is also related to human activity such as intentional or accidental introduction, spread by ballast waters and underwater constructs of vessels, etc.

Four of the introduced fish species with unindentified vector of appearance (mediterranisation or spread by ballast waters) belong to the benthic ecological group that lay their eggs and have their offspring at the bottom. The spread of such species with ballast waters is almost impossible since ballast tanks are filled above a considerable depth to avoid clogging. In such case, these species with a high probability can be included into the group of fishes that have spread naturally (mediterranisation).

Noteworthy that the increase in the number of introduced fish species in waters of Crimea in the past 50 years is also a result of detailed monitoring of the regional ichthyofauna, which has been carried out by researchers of A. O. Kovalevsky Institute of Biology of Southern Seas NAS of Ukraine using modern trapping tools.

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