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Reproduction and Life History Strategies of the Common Jellyfish

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Reproduction and Life History Strategies of the Common Jellyfish Hydrobiologia 451: 229–246, 2001. 229 © 2001 Kluwer Academic Publishers. Printed in the Netherlands. Reproduction and life history strategies of the common jellyfish, Aurelia aurita, in relation to its ambient environment Cathy H. Lucas School of Ocean & Earth Science, University of Southampton, Southampton Oceanography Centre, European Way, Southampton, SO14 3ZH, U.K. Tel: +44 (0)23 8059 6270. Fax: +44 (0)23 8059 3059. E-mail: [email protected] Key words: medusa, polyp, reproduction, recruitment, temperature, food Abstract The scyphozoan Aurelia aurita (Linnaeus) is a cosmopolitan species, having been reported from a variety of coastal and shelf sea environments around the world. It has been extensively studied over the last 100 years or so, and examination of the literature reveals three striking features: (1) the presence of populations in a wide range of environmental conditions; (2) large inter-population differences in abundance and life history patterns over large and small spatial scales; and (3) inter-annual variability in various aspects of its population dynamics. A. aurita is clearly a highly flexible species that can adapt to a wide range of environmental conditions. While various physiological and behavioural characteristics explain how A. aurita populations can take advantage of their surrounding environment, they do not explain what governs the observed temporal and spatial patterns of abundance, and the longevity or lifespan of populations. Understanding these features is necessary to predict how bloom populations might form. In a given habitat, the distribution and abundance of benthic marine invertebrates have been found to be maintained by four factors: larval recruitment (sexual reproduction), migration, mortality and asexual reproduction. The aims of this review are to determine the role of reproduction and life history strategies of the benthic and pelagic phases of A. aurita in governing populations of medusae, with special attention given to the dynamic interaction between A. aurita and its surrounding physical and biological environment. Aurelia aurita populations between an asexual benthic polyp, or scyphistoma, and a sexual pelagic medusae. Examination of the General observations wealth of literature on A. aurita populations reveals three striking features, summarised as follows: (1) The common moon jellyfish, Aurelia aurita (Lin- the presence of populations in a wide range of en- naeus), is a cosmopolitan species with a worldwide ◦ ◦ vironmental conditions; (2) large inter-population dif- distribution in neritic waters between 70 N and 40 S ferences in abundance and life history patterns, and (Kramp, 1961; Russell, 1970). It has been reported (3) inter-annual variability in abundance, growth, and from a variety of coastal and shelf sea marine envir- timing of reproduction within single populations. onments particularly in northwestern Europe (Palmén, 1954; Möller, 1980; Hernroth & Gröndahl, 1983; Van Habitats Der Veer & Oorthuysen, 1985; Lucas & Williams, 1994; Olesen et al., 1994; Schneider & Behrends, A. aurita populations are found predominantly in 1994), the Black Sea (Shushkina & Arnotauv, 1985; coastal embayments, fjords, and estuaries where there Lebedeva & Shushkina, 1991; Mutlu et al., 1994), Ja- are suitable substrata for the benthic scyphistoma pan (Yasuda, 1969; Ishii et al., 1995; Omori et al., polyp. Within these habitats, the degree of contain- 1995), and parts of North America (Hamner et al., ment, tidal flow, water depth, temperature and salinity, 1994; Greenberg et al., 1996). A. aurita has a com- and trophic condition can vary quite considerably plex life cycle, comprising an alteration of generations (Table 1). 230 Table 1. Summary of the documented habitats of Aurelia aurita medusae, together with their maximum abundance and biomass. Location Types: s-e est = semi-enclosed estuary; s-e l = semi-enclosed lagoon; s-e fjd = semi-enclosed fjord; o-fjd = open fjord; o-b = open bay; s-e b = semi-enclosed bay; o = open water. nd = no data. aConverted from published data using WW:DW:C relationships reviewed by Hirst & Lucas (1998). References: 1. Lucas & Williams (1994); 2. Lucas (1996); 3. Olesen et al. (1994); 4. Riisgård et al. (1995); 5. Hernroth & Gröndahl (1983); 6. Gröndahl (1988a); 7. Omori et al. (1995); 8. Panayotidis et al. (1988); 9. Hamner et al. (1982); 10. Ishii & Båmstedt (1998); 11. Schneider (1989b); 12. Möller (1980); 13. Schneider & Behrends (1994); 14. Mutlu et al. (1994); 15. Van Der Veer & Oorthuysen (1985); 16. Yasuda (1969); 17. Yasuda (1971); 18. Rasmussen (1973) Location Type Depth Notes Productivity Max. Max. Ref (m) Abundance Biomass − − (No. m 3)(mgCm3) − Southampton Water, U.K. s-e est 15 industrial High (12.8 mg C m 3 mesozooplankton) 8.71 30.2 1,2 − Horsea Lake, U.K. s-e-l 6–7 no river Low (5.2 mg C m 3 mesozooplankton) 24.9 43.2 2 Kertinge Nor, Denmark s-e fjd 2–3 Eutrophic + low mesozooplankton 300 350–450 3,4 Gullmarfjord, Sweden 0-fjd 120 winter ice nd 14.96 nd 5,6 − Tokyo Bay, Japan o-b nd Eutrophic (400 mg DW m 3 zooplankton) 1.53 125a 7 − Elefsis Bay, Greece s-e b 33 anoxia Eutrophic (471 mg DW m 3 zooplankton) 44 95a 7 Jellyfish Lake, Palau s-e l 30 sulphide High 0.30 nd 9 Vågsbøpollen, Norway s-e b 12 Very low zooplankton 22.3 710 10 − Eckernförde Bay, Kiel Bight o-fjd 20 26.7 mg C m 3 zooplankton 0.03–0.23 54.8 11 Kiel fjord, Kiel Bight o-fjd 8 nd 0.33 ∼60 12,13 Kiel Bight o nd nd ∼0.14 ∼35 13 − Black Sea o >200 Mneiopsis Eutrophic ? 0.21 g C m 2 14 Dutch Wadden Sea o 5–10 North Sea High ? 0.49 175 15 Urazoko Bay, Japan s-e b nd nd 71 nd 16,17 Isefjord, Denmark o-fjd 9 winter ice Eutrophic nd nd 18 Medusa abundance is generally higher in small, shallow, semi-enclosed or enclosed systems with lim- ited tidal exchange (Olesen et al., 1994; Lucas, 1996; Ishii & Båmstedt, 1998), than in open water sys- tems or where depths exceed several hundreds of metres (Van Der Veer & Oorthuysen, 1985; Mutlu et al., 1994; Schneider & Behrends, 1994). On a seasonal and small-scale spatial basis, temperate es- tuaries often have widely fluctuating temperature and salinity regimes, and A. aurita can be considered both eurythermal and euryhaline in its distribution (Fig. 1). Populations occur in fjords that experience winter ice cover (Rasmussen, 1973; Hernroth & Gröndahl, 1983, 1985a), and in tropical, aseasonal environments such as Jellyfish Lake, Palau, where surface water temper- ature rarely deviates from 31 to 32 ◦C (Hamner et al., 1982; Dawson & Martin, 2001). Similarly, A. aur- ita are found in salinities ranging from 14 to 22 in Kertinge Nor, Denmark (Olesen et al., 1994) to 38 in Elefsis Bay, Greece (Papathanassiou et al., 1987), although there are also reports of populations living in salinities of <10 (see Russell, 1970). As well as the physical parameters described Figure 1. Temperature and salinity ranges of Aurelia aurita. above, one of the most conspicuous aspects of A. 231 aurita distribution is its presence in systems with Jellyfish Lake (Hamner et al., 1982) and Horsea Lake highly contrasting nutrient inputs, productivity and (Lucas, 1996). food availability (Table 1). For example, Horsea Lake Patterns of growth and the maximum bell diameter in southern England has no riverine input to sup- attained by medusae are highly variable among pop- ply nutrients, and a numerically- and species-poor ulations. Three growth phases are typical of most zooplankton community that is controlled by both populations: slow growth during the winter and early bottom-up (chlorophyll a <2mgm−3) and top-down spring; exponential growth once temperature and food (A. aurita 43 mg C m−3) pressure (Lucas et al., 1997). availability have increased in mid-spring; and finally, On the other hand, A. aurita populations may also shrinkage in the summer and autumn during gam- be abundant in polluted, eutrophic systems that may ete release (Möller, 1980; Schneider, 1989b; Lucas have anoxic bottom waters. Examples include Elefsis & Williams, 1994). Medusae typically reach 200– Bay (Papathanassiou et al., 1987; Panayotidis et al., 300 mm maximum bell diameter, although they can be 1988), Kertinge Nor (Olesen et al., 1994; Riisgård et as large as 450 mm (M. Omori, pers. comm.). Com- al., 1995) and the Black Sea (Mutlu et al., 1994). parison among populations reveals an inverse relation- ship between medusa abundance and maximum bell diameter (Fig. 3), suggesting that there is a density- Variations among populations dependent mechanism governing A. aurita popula- tions, as was first suggested by Schneider & Behrends A. aurita populations around the world are character- (1994) for the Kiel Bight population. In severely food- ised by a great diversity in population and life history limited populations such as in Kertinge Nor (Olesen characteristics, such as abundance, growth, timing and et al., 1994) and Horsea Lake (Lucas, 1996), instant- − periodicity of strobilation, timing of and size at sexual aneous growth rates are <0.1 d 1, and mean bell maturation and longevity of the medusa (Fig. 2). Sig- diameter of adult medusae is typically <50 mm. nificant differences in population characteristics can Medusae typically reach sexual maturation at the occur over small (10s of kilometres) and large (100s upper end of the size range of the population (Lucas of kilometres) spatial scales, with the role of envir- & Lawes, 1998). In most situations adult medusae onmental conditions such as temperature and food shrink and die following spawning. It is thought that availability considered to play a key role. this is caused by extrusion of gastric filaments during In many of the populations studied, the main gamete release, resulting in morphological degrada- period of strobilation, resulting in the liberation of tion and susceptibility to parasitic invasion (Spangen- ephyrae, starts in the late winter/early spring (Thiel, berg, 1965).
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