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On the Fisheries of the Black Sea and Caspian Sea Impacts Of BLACK SEA OCEANOGRAPHY Impacts of Invasive Ctenophores on the Fisheries of the Black Sea and Caspian Sea Figure 1b. Th e ctenophore Beroe ovata of the Black Sea. Th is specimen has no food Figure 1a. Th e ctenophore Mnemiopsis leidyi of (i.e., Mnemiopsis leidyi) in its stomach, which is most of the body cavity (compare the Black Sea. Photo by Ahmet E. Kideys. with Figure 1c and 1d where the organisms have food in their stomachs). Mouth of B. ovata, which is half closed, is along the entire width. Th is article has been published in Oceanography, Volume 18, Number 2, a quarterly journal of Th e Oceanography Society. Copyright 2005 by Th e Oceanography Society. All rights reserved. Reproduction of any portion of this article by photo- 76 Oceanography Vol.18, No.2, June 2005 copy machine, reposting, or other means without prior authorization of Th e Oceanography Society is strictly prohibited. Send all correspondence to: [email protected] or Th e Oceanography Society, PO Box 1931, Rockville, MD 20849-1931, USA. BY AHMET ERKAN KIDEYS, ABOLGHASEEM ROOHI, SIAMAK BAGHERI, GALINA FINENKO, AND LYUDMILA KAMBURSKA Gelatinous zooplankton are known to have a substantial effect on pelagic food webs by exerting top- down control on their ecosystems (Purcell and Decker, 2005), but their impact could be even greater when they invade other regions. The effects of zooplankton invasions have been well documented for the Black Sea and very recently for the Caspian Sea. Until the 1980s, three species of gelatinous macrozooplankton were present in the Black Sea: two scyphozoan medusae (Rhizostoma pulmo and Aurelia aurita) and only a single ctenophore species (Pleurobrachia pileus) (Kideys and Romanova, 2001). Then, in the early 1980s, the ctenophore Mne- miopsis leidyi (Figure 1a), already known as a voracious zooplankton feeder (Reeve et al., 1978), was transported (likely via ballast waters) to the Black Sea from its native waters of the western Atlantic Ocean. Achievement of peak biomass levels of this ctenophore at the end of the 1980s caused the onset of striking adverse ecological events in the Black Sea, such as abnormal oscillations in plankton dy- namics (Kideys et al., 2000) and, most notably, a sharp decrease in the pelagic fi shery (Kideys, 1994). By the end of the 1990s, the accidental introduction of another ctenophore to the Black Sea, Beroe ova- ta (already known to feed exclusively on other ctenophores) (Figure 1b, 1c, 1d), coincided with another set of striking events, but this time it was benefi cial for the ecosystem and the fi shery. Figure 1c. Th e ctenophore Beroe ovata of the Black Sea with a Figure 1d. Th e ctenophore Beroe ovata just swallowed a Mnemiopsis half-digested Mnemiopsis leidyi specimen in its stomach. leidyi specimen in the Black Sea. Photo by Ahmet E. Kideys. Oceanography Vol.18, No.2, June 2005 77 Invasion of these ctenophores and et al., 1978). Such high growth rates can et al., in press). M. leidyi has also been their contribution to the decline of the only be achieved with a very high feeding shown to feed on the eggs and larvae of entire Black Sea’s ecosystem in general, rate. The abundance of Mnemiopsis has fi shes both in its native region and in the and on its fi shery in particular, has been been observed to fl uctuate throughout Black Sea (Mutlu, 1999). questioned. Other factors, such as over- the year. In the Narragansett Bay (off- In the Black Sea, M. leidyi (as well as fi shing (Gücü and Oguz, 1998; Gücü, shore Rhode Island, USA), for example, Aurelia aurita and Beroe ovata) are con- 2002) and climate change (see Oguz, this Kremer and Nixon (1976) found that fi ned to surface (and mainly coastal) issue) may also be to blame. All of these while the overwintering population of waters above the thermocline (in the factors could be acting simultaneously Mnemiopsis was extremely low (1 to 2 warm seasons) (Mutlu, 1999; Kideys and on the ecosystem, however, pinpoint- animals per 104 m3), they could reach a Romanova, 2001) coincident with the ing the major ones is very important for peak density of over 50 individuals per greatest abundance of zooplankton (due understanding sensitive ecosystems that m3 in summer. Although highest abun- to smaller copepod and cladoceran spe- are increasingly exposed to anthropo- dances occur in summer (and early au- cies as well as young stages of other im- genic effects and for using the experience tumn), unlike in native regions, relatively portant forage zooplankton). These ge- obtained from the Black Sea case study high concentrations of this ctenophore latinous zooplankton are separated from to predict, and possibly mitigate, future were present during other seasons in the the native ctenophore Pleurobrachia rho- ecosystem decline. One immediate prac- Black Sea (Mutlu, 1999; Kideys and Ro- dopis, which dwells in deeper (below the tical benefi t of such knowledge would manova, 2001). It is worth noting that thermocline at around 30 m in summer) be for helping to stem the decline of the Mnemiopsis’s main summer spawning and offshore waters. neighboring Caspian Sea, a unique and period coincides with that of the ancho- fertile ecosystem, which is presently ex- vy in the Black Sea. LONGTERM DISTRIBUTION OF periencing similar adverse events, but on Mnemiopsis has long been reported M. LEIDYI IN THE BLACK SEA an even greater scale. as an effective predator on zooplankton. The biomass of the offshore- (and deep-) Reeve et al. (1978) state that perhaps dwelling ctenophore Pleurobrachia rho- BIOLOGY OF MNEMIOPSIS the most signifi cant aspect of the feed- dopis varies less over time (Kideys and LEIDYI IN ITS NATIVE AND ing behavior of Mnemiopsis is that their Romanova, 2001) than Aurelia aurita INTRODUCED REGIONS ingestion rate is proportional to food and M. leidyi. Figure 2 shows long-term Mnemiopsis (Ctenophora) has certain concentration. Because of this, concomi- changes in the biomass of only M. leidyi biological characteristics that enable this nant with their appearance, a dramatic and Aurelia aurita (biomass changes of species to signifi cantly affect its ecosys- decrease in the number or biomass of the most recent ctenophore Beroe ovata tem. Main (1928 cited in Vinogradov et copepods and other food zooplankton is are discussed separately later). In this al., 1989) points out that, although most often observed in the native region. For fi gure, the values obtained until 1991 medusae are microphages (consume mi- example, Burrell (1968, cited in Kremer, croscopic prey), Mnemiopsis is a macro- 1979) suggested that predation by M. Ahmet Erkan Kideys ([email protected]. phage capable of consuming fairly large leidyi was responsible for a large frac- edu.tr) is Associate Professor, Institute of prey (up to 1 cm or more in length). tion (73 percent) of total zooplankton Marine Sciences, Erdemli, Turkey. Abol- Mnemiopsis, like all other ctenophores, mortality in the York River estuary of ghaseem Roohi is Research Associate, are hermaphroditic. Ctenophores, partic- the Chesapeake Bay (offshore Virginia, Mazandaran Fisheries Research Center, Sari, ularly members of the genus Mnemiopsis, USA). Finenko and Romanova (2000) Iran. Siamak Bagheri is Research Associate, have a very high reproductive capacity. calculated that on a daily basis, about Guilan Fisheries Research Center, Anzali, Mnemiopsis mccradyi is able to produce 20 percent of total forage zooplankton Iran. Galina Finenko is Senior Research 8,000 eggs within 23 days—after only 13 was being eaten by this ctenophore in Associate, Institute of Biology of the South- days of its own birth. The growth rate the summer of 1995. Similar high values ern Seas, Sevastopol, Ukraine. Lyudmila of this species is comparable to that of were found by other scientists studying Kamburska is Senior Research Associate, phytoplankton (daily doublings) (Reeve the Black Sea and Caspian Sea (Finenko Institute of Oceanology, Varna, Bulgaria. 78 Oceanography Vol.18, No.2, June 2005 25 belong to Russian investigators (mainly 25 (a) Bulgaria 20 20 from the western and northern Black 15 15 Sea), and after 1991, to Turkish investi- 10 10 gators (from the southern Black Sea). It 5 5 should be noted that, although the Turk- 0 0 1970 1975 1980 1985 1990 1995 2000 ish data were from a higher number of 25 25 (b) Romania stations, Kideys and Romanova (2001) 20 20 ) -2 suggested these values were too low be- 15 15 cause their nets were not as effi cient (had 10 10 a coarser mesh) as the Russians’. Because 5 5 of this, the Russian values were effi cien- 0 0 cy-corrected (multiplied by 2 to 3 based 1970 1975 1980 1985 1990 1995 2000 on the net most often used), though 25 500 (c) Former USSR Countries those from the southern Black Sea (af- 20 400 ter 1991) were reported as uncorrected, 15 300 fish catch (thousand tons) Total 200 raw values. As seen in Figure 2, after the 10 Forage zooplankton biomass (g wet wtForage m 5 100 famous peak values of 1989, biomass 0 0 values for M. leidyi as high as 485±34 g 1970 1975 1980 1985 1990 1995 2000 2 25 WW/m (wet weight per meters squared) (d) Turkey 500 20 were obtained in 1995 (uncorrected, raw 400 15 300 data from March, which is not the main 10 200 development season of this ctenophore!) 5 100 from a dense station grid (107 stations) 0 0 1970 1975 1980 1985 1990 1995 2000 in the southern Black Sea (Mutlu, 1999). 2000 . 2000 1500 These data clearly show that the Rus- 1600 1430 1200 900 biomass sian (corrected) values of 1.5 to 2 kg ) 800 -2 WW/m2 obtained for early autumn 1989 Mnemiopsis leidyi (Vinogradov et al., 1989) are reasonable aurita A.
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