Shads of Eastern Europe from the Black Sea: Review of Species and Fisheries
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American Fisheries Society Symposium 35:69-7 6, 2003 O 2003 by the American Fisheries Society Shads of Eastern Europe from the Black Sea: Review of Species and Fisheries IoN Navooenur DantLbe Deltn National Instítute for Reseatch and Deaelopntertt, Babadng Street, 165 Tilcea 8800, Romanin JouN R. Walpr,reN HtLdson Rioer Foundntion for Science nnd En-oironmental Research, 77 Bnttery Place, Suite 915, Neu York, Netn York 10004, USA Abstrnct species beÌonging to Alosa are known in the northwestern part of the BÌack -Three Sea, including the Sea of Azov: Pontic shad A. pontica (EichwaÌd, 1838), Caspian shad Á. caspia (Etchwald, 1838), and BÌack Sea shad A. mneoticn (Crimm, 1901). Regional populations, and possibly subspecies, occur in the Danube, Dniester, Dnieper, and Don rr'vers. Pontic shad, the most abundant shad of Eastern Europe, migrates 500 km upstream for spawning in the Danube River at the end of March in waters oÍ 3-5'C. The peak of these movements occursinApril May at9-TT"C,withtherunendinginJuly-Augustal22-26"C. Pelagiceggs and larvae drift seaward in the upper 0-0.5-m layer of river water. Young-of-the-year Pontic shad feed in brackish waters near the Danube delta, with juveniles leaving the shoreline in autumn. The adult stock lives at depths oÍ 50-100 m, Íeeding mainly on fish. The longevity of Pontic shad is 5-7 years, with maturity occurring at 3-4 years. Most die after the spawning migration, with less than 10% spawning repeatedly. The other two shad species spawn in brackish water of deltas and lagoon Ìakes. The Pontic shad fishery is the most important, followed by that of Caspian shad; the Black Sea shad fishery has collapsed. Fishing Pressure and pollution are the main threats for Pontic shad. However, spawning habitat Íor Caspian shad and Black Sea shad have drastically decreased due to damming. River fisheries for Pontic shad are managed primarily through season and atea closures. Species behavior, spawning and recÍuitment, year-class strength, and estimation of popuÌation dynamics remain open research and management issues. lntroduction The Black Sea Three alosines are native to Eastern Europe: Pontic The Black Sea is a large inland sea measuringT,2l1 shad Alosa pontica (Eichwald, 1838), Caspian shad km from east to west and as much as 560 km from A. caspia nordmanni (Eichwald, 1838), and Black Sea north to south, with an area of more than 400,000 shad Á. mneotica (Grimm, 1901). These shads are km2. It is bordered by Ukraine on the north, Russia heavily Íished and are important commercially and on the northeast, Georgia on the east, Turkey on culturally within their respective distributions. All the south, and Bulgaria and Romania on the west. are found in the Black Sea region, an area of con- The Black Sea is connected with the Mediterranean siderabÌe environmental stress. However, knowl- Sea by a reÌatively narrow water\ rav that includes edge of the biology and conservation status of these the Bosporus, the Sea of Marmara, and the shads is poor. Dardanelles. Because of this, its tides are generalÌy In this paper, we review the biology and sta- less than 10 cm. The Black Sea's largest arm is the tus of all three shads, with an emphasis on the best- Sea of Azov; both the Don and Kuban rivers enter known species, Pontic shad. Both Pontic shad and it. Other major rivers flor.'r'ing into the Black Sea Caspian shad are found in the Black Sea and include the Southern Buh, Danube, Dneister, and Caspian Sea basins; we restrict our treatment to the Dnieper. About 20% of Europe's continental land- BÌack Sea basin. Coverage of these fishes in the mass drains to the Black Sea. Caspian Sea is provided in Coad et al. (2003, this The Black Sea has two lavers of water of dif- volume). ferent densities. The bottom layer is highly saline, rich with hydrogen suÌfide, and hypoxic, has lìttle Ì Corresponding author: [email protected] circulation, and is virtually dead biologicalÌy. The 69 lsu Welou'qN 70 Nevooanu countercloc'k- laver is less saline' circulates 'nner lire' However' the ì:f5 ";à';";À,i' ,i.n marine ;h.k;;ã,-'iilto"*""thasbeent"";l"t-t"tffo"lï tive sPecies, blue' or viol is leidYi (also flanks' TheY the oPercuÌa bu ected. wj.th the the oPerculum' origin accoun distinguish mon Ponto-Caspian Thethreespe1L^ +Lrrêê qnê .tntiçn:t:;to u*o"g theiifaunas raker counts are useful ;Ë;;"d where they co-oc Gill ad'ults but not for discriminatory fe es among TaxonomY Young sPecimens cm. nlosn wlth Alosn' Banarescu (1964) Distribution shad in the Black signations have Pontic Shad ludes the :1ïi:ïi:.fl-"ï:"Ï il: of Azov to sPawn e.been dis- srdes of the between astern PoPulation ranges Crimean ch near túe southeastern A. tnnaícn, to be a single A. tnnnicn has PriorìtY shouÌd be considered va Pontic shad and Bl ern Crimean Peninsula' on both jaws are well deveioPe tic shad and Black Sea shad a ìn CasPian shad' In all ering ierminates with alae, which iie sYmme caudaÌ fin The ventra 77 Snnos op E,qsreRN Eunopp FRoM rHE Blecr Sre :ends along the Rom Dur- BiologY jìg some years, it is Lake ìn spawning :nd the Razim-Sinoi escu- There are considerable differences eonte and Munteanu 1968). During spring, it is ;r idespread in the Danube delta and adiacent flood- :lain lakes (Antipa 1905; Leonte 1943; Banarescu r964). Some individuals have been found 375 km Kolarov 1991b) and 496ktn (Kovachev 1922) :up- is known of the spawn- :tream from the Danube River mouth. Berg (1948) rivers or in river deltas' Less shad, but it appears they arrd Svetovidov (1952, 1964) cited Antipa's (1905, ing iocations of Black Sea of deltas and lagoons 1909) opinion that Caspian shad migrate up to km utilize low-salinity waters (1985) discussed two 931 (Iron Gate) from the Danube's mouth, and this Ivanov and Beverton in the ivas recently corroborated by N. Bacalbasa- Íorms of Pontic shad. The larger one spawns part of Dobrovici (Lower Danube University, personal Don River and overwinters in the eastern and Batumi' communication), who caught Caspian shad at river the Black Sea between Novorossisk in the western portion of km 864. Caspian shad is also found in the Dniester The smaller form lives the Danube River to and Bug rivers, but it never migrates far from their the Black Sea and enters is high, it aiso river mouths. In the Dnieper Riveç it does not mi- ance n (Vladimirov 1953a). The r, and Bug rivers. had Á. c. tanaica extends migrated in the 90 Sea, north of Batumi, the Dnieper River to Kiev but now can travel only mouth due to a dam Sea of Azov and the Lower Don River. km from the river ' Pontic shad migrate northward in the Black Black Sea Shad Sea in spring and early summer as they move to- rivers. Historically, Pontic shad This species inhabits (Figure 1) both the Black Sea ward spawning penetrated far up the Danube with specimens cap- and Sãa of Azov (Kolarov 1991c). Large schools of at Budapest (river km 1,650), but the migra- this fish enter in Razim-sinoie Ìake complex on the iured route was reduced to 864 km, aÍter construc- Romanian coast (Borcea 1936; Teodorescu-Leonte tion of the Iron Gate II hydroelectric plant in 1983' and Munteanu 1968). It also appears in some years tion Barrages at river km 943 and 864 appear not to along the Bulgarian coast. In the eastern part of the have Àad a large impact on the river's Pontic shad Blaci Sea, it ranges from the region of the Strait of occurs between river Kerch to Novorossysk and south along the popuiation; most spawning and 500. Caucasus coast. km 180 rrì ìlâ ,| tl \ ii Ìì'1 ag,.1!'ar, -:-'€.\., ' l .- tl .f - tlümáfi1á 6!tl:.- *' I ; I I s aí'ii Elack5ea Trrlêy I O Black Sea shad, and (c) Caspìan shad' from the Figure 1-Distribution of Eastern European shads, (a) Pontic shad, ft) Black Sea. 72 Navooenu aNIo WeloMAN Adult Pontic shad moving upriver in the tic shad entering the Danube River was strongly Danube for spar,r'ning are composed of age-2 to age- positive (y = 0.4832x - 7 .4397 ; r' = 0.9764), possibly 7 individuaÌs. These proportions shift annualÌy, but compensating for loss of biomass through greater age-3 specimens usually comprise 50ok or more of energy expenditure in migration and reproduction the totaÌ run, with the age-6 and age-7 classes con- at higher water temperatures. There was also a tributing about 1% or less (Navodaru 7996, 1998). strongly positive relationshrp (rt = 0.8551) between The three shad species have similar maximum river migration distance at capture and body con- kno'wn ages: 6 years for Black Sea shad and 7 years dition as assessed by the Fulton index (Figure 3). for the other two species. During this upstream migration, Ponüc shad / Total lengths oÍ Pontic shad males and females spawners lost biomass due to egg maturation and ; are approximately equal for each age-class; femaÌes deposition and energy spent for migration. Analy- r,r'eigh more than males at each age interval, but this sis of nine postspawning individuats in compari- difference is not statistically significant (P < 0.05). son with their biomass entering in-river, calculated The relationship between total length (TL) and to- with the length-weight relationship, indicated that taÌ weight (TW; TW = aTLb), estimated for 6 years these shad iost of 10.3-42.7% of biomass.