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Trophic Dynamics of Two Sympatric Mysid Species in an Estuarine Transition Zone

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Trophic Dynamics of Two Sympatric Mysid Species in an Estuarine Transition Zone MARINE ECOLOGY PROGRESS SERIES Vol. 332: 171–187, 2007 Published March 5 Mar Ecol Prog Ser Trophic dynamics of two sympatric mysid species in an estuarine transition zone Gesche Winkler*, Christine Martineau, Julian J. Dodson, Warwick F. Vincent, Ladd E. Johnson Québec-Océan, Département de biologie, Université Laval, Ste-Foy, Quebec G1K 7P4, Canada ABSTRACT: The feeding ecology of 2 sympatric mysid species within the food web of the St. Lawrence Middle Estuary was studied to determine their trophic position and potential mechanisms of coexistence. These abundant predators were clearly distinguished by differences in foraging behaviour. In feeding experiments involving multi-prey assemblages, predation and ingestion rates of Mysis stenolepis were greater than for Neomysis americana, except for small prey. M. stenolepis showed highest predation rates on and preference for Eurytemora affinis nauplii and copepodites, and did not feed on the most abundant prey, the veliger stage of the zebra mussel Dreissena poly- morpha. In contrast, N. americana showed highest predation rates on and preference for small prey species such as rotifers, nauplii and veligers. When densities of preferred prey declined in the second half of the experiment, both mysid species showed a shift in predation towards the less preferred prey. M. stenolepis switched to filter feeding, with higher predation rates on veligers, whereas N. americana increased predation on small E. affinis copepodites. Stable isotope analysis and feeding experiments suggested 3 trophic levels in the food web, with a high degree of omnivory. The δ13C values for the 2 mysid species were similar and supported by the same carbon source, mostly of autochthonous origin. However, their δ15N differed significantly by 2.2‰, indicating that the compo- sition of their diet differed in terms of ingested and assimilated food items. These results suggest that co-existence of the sympatric mysid populations is achieved by reduced competition due to food- niche partitioning. Flexibility in their foraging behaviour allowed mysids to exploit multiple prey groups that undergo temporal and spatial fluctuations in the highly dynamic estuarine transition zone of the St. Lawrence Middle Estuary. KEY WORDS: Mysis stenolepis · Neomysis americana · Feeding experiments · Zooplankton assemblage · Stable isotopes Resale or republication not permitted without written consent of the publisher INTRODUCTION impact on zooplankton community structure has been well studied in lakes, where mysids have been intro- Mysids are a major component of estuarine and duced to improve salmonid fisheries, resulting in major coastal zooplankton communities in terms of both declines in the abundance of cladocerans and, at times, abundance and biomass. Due to their importance as a copepods (Morgan 1981, Langeland 1988). In estuar- food source for fishes, there are many studies of their ies, high mortality rates of copepods have been distribution and population dynamics (for review see observed, due to the predation pressure of mysids Tattersall & Tattersall 1951, Mauchline 1980), but their (Johnston & Lasenby 1981, Jerling & Wooldridge importance in trophic dynamics is still not well under- 1995). stood. Mysids are generally considered to be omni- Two sympatric mysid species, Mysis stenolepis and vores, and adjust their foraging behaviour according to Neomysis americana, occur in many estuaries of north- food availability (Mauchline 1980). Their potential eastern North America, including the St. Lawrence *Email: [email protected] © Inter-Research 2007 · www.int-res.com 172 Mar Ecol Prog Ser 332: 171–187, 2007 Estuary. The estuarine transition zone (ETZ) of the St. cally involve 1 or 2 prey (Fulton 1982, Gorokhova & Lawrence Middle Estuary is a highly productive region Hansson 1997, Viitasalo & Rautio 1998, Viherluoto & for both mysid species. The ETZ is characterised by Viitasalo 2001a), which severely limits interpretation sharp gradients in salinity, temperature and nutrient within a complex food web. Interactions among differ- content, and hydrodynamic forces create strong estu- ent prey types as a function of their abundance, size arine re-circulation. This acts as a retention mecha- and escape behaviour may play an important role in nism for inorganic and organic matter, resulting in determining the predation rate on each individual high turbidity and high biomass of primary producers, species in their natural environment. For example, herbivores and planktivores (Frenette et al. 1995, Vin- predation rates on copepod Acartia tonsa by Praunus cent et al. 1996, Winkler et al. 2003). The zooplankton flexuosus were reduced when a second copepod prey, community in the transition zone is composed of 3 Eurytemora affinis, was present (Viitasalo & Rautio distinct assemblages that are mainly determined by 1998). average salinity and the degree of water column strat- Short-term experiments can only provide infor- ification: (1) a tidal-freshwater assemblage in waters of mation on potential trophic interactions. Fortunately, salinities <1.6 psu, characterised by high abundances stable isotope analysis can provide integrated esti- of veligers of the bivalve invader Dreissena poly- mates of carbon source trophic position. This tech- morpha, rotifers, the amphipod Gammarus tigrinus nique gives a time-integrated view of the food assimi- and cladocerans (mostly Bosmina longirostris); (2) an lated by an organism, accounting for complex food estuarine assemblage in weakly stratified waters of web interactions such as omnivory (Cabana & Ras- salinities between 0.5 and 6 psu, characterised by mussen 1996). Stable carbon isotope ratios (δ13C) abundance peaks of the calanoid copepod Eurytemora reveal the nature of the carbon source ingested, as the affinis and 2 mysid species M. stenolepis and N. amer- signature increases only slightly in the consumer dur- icana; and (3) a euryhaline marine assemblage com- ing assimilation of food sources (Peterson & Fry 1987, posed of high abundances of marine copepods such as France & Peter 1997, Johannsson et al. 2001). How- Calanus spp. and Acartia spp. (Laprise & Dodson 1994, ever, the stable nitrogen isotope ratios (δ15N) can be Winkler et al. 2003, 2005). High abundances of M. used to identify trophic position (Cabana & Rasmussen stenolepis and N. americana are found during the sum- 1996), as the signature of a consumer is typically mer months when zooplankton standing stock is also enriched by 3.4(±1)‰ relative to its food source (Mini- high (G. Winkler, L. E. Johnson, J. J. Dodson unpubl. gawa & Wada 1984). Recent work on the diet of Mysis data). Little is known about the trophic dynamics, relicta with stable isotope signatures in combination niche overlap and potential competition of these sym- with stomach content analysis showed more infor- patric species in the food web of the ETZ. mative results (Johannsson et al. 2001) than each The feeding ecology of mysids has traditionally been approach by itself. Stomach content analysis identified assessed by the analysis of stomach contents (Gross- diatoms as an important prey item, but only with stable nickle 1979, Fockedey & Mees 1999, Viherluoto et al. isotope analysis were those authors able to show that 2000). Stomach content analysis of both mysid species diatoms accounted for 50% of the energy intake of the Mysis stenolepis and Neomysis americana in the ETZ mysids. revealed a diverse diet of calanoid and harpacticoid The purpose of this study was to determine the copepods, cladocerans, rotifers, diatoms and pollen, trophic position of the mysid species Neomysis ameri- suggesting omnivorous foraging behaviour (Winkler et cana and Mysis stenolepis in the food web of the estu- al. 2003). Feeding rates, preferences and selection of arine transition zone of the St. Lawrence Estuary. We these food items are not known. Elsewhere, N. ameri- also examined the potential for competitive inter- cana has been shown in experiments to prey on several actions between these 2 species, leading to resource marine copepods such as Acartia tonsa and Centro- partitioning. To address these objectives, we carried pages spp., although gut content analyses revealed a out laboratory feeding experiments with naturally oc- dominance of detritus (Fulton 1982). This approach curring zooplankton assemblages to include the poten- has its limitations: stomach content analysis shows tial of interference among different prey species, while ingested food and the relative proportion of each item quantitatively evaluating predation rates, prey pre- ingested, but underestimates easily digested items, ference and prey selection by N. americana and M. such as soft-bodied rotifers, and no estimates of inges- stenolepis. In addition, we compared the experimental tion rates are possible. Ingestion rates can, however, results with the stable isotope signatures of both mysid be assessed by feeding experiments designed to define species and their potential mesozooplankton prey in functional responses to predator size, prey size and the ETZ to assess quantitatively the role of mysid prey density (Webb et al. 1987, Wooldridge & Webb species in energy transfer and their impact on prey 1988, Winkler & Greve 2004), but these studies typi- communities. Winkler et al.: Trophic dynamics of two sympatric mysid species 173 MATERIALS AND METHODS the zooplankton sample to calculate prey concentra- tions that we introduced into the experiments. We Predation experiments. Prey selection and diet
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