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Cherel, Yves, Camille Fontaine, Pierre Richard, and Jean-Philippe Labat

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Cherel, Yves, Camille Fontaine, Pierre Richard, and Jean-Philippe Labat Limnol. Oceanogr., 55(1), 2010, 324–332 E 2010, by the American Society of Limnology and Oceanography, Inc. Isotopic niches and trophic levels of myctophid fishes and their predators in the Southern Ocean Yves Cherel,a,* Camille Fontaine,a Pierre Richard,b and Jean-Philippe Labatc a Centre d’Etudes Biologiques de Chize´, Centre National de la Recherche Scientifique, Villiers-en-Bois, France bLittoral, Environnement et Socie´te´s, Centre National de la Recherche Scientifique et Universite´ de La Rochelle, La Rochelle, France c Universite´ Pierre et Marie Curie-Paris 6, UMR 7093, LOV, Observatoire Oce´anologique, Villefranche-sur-Mer, France Abstract We report the trophic structure of a myctophid assemblage by measuring the isotopic niches of 14 species living in Kerguelen waters, southern Indian Ocean. Most of the species show distinct isotopic niches that differ by at least one of the two niche axes (d13C habitat and d15N trophic position), indicating trophic partitioning within the assemblage. Strong niche segregation occurs within each of the three most common genera of myctophids (Electrona, Gymnoscopelus, and Protomyctophum), illustrating the different mechanisms (habitat and dietary segregation) that allow coexistence of closely related species. Calculated trophic levels (TLs) of myctophids ranged from 3.3 to 4.2, showing that they are secondary and tertiary consumers in the pelagic ecosystem. The positive relationship between TL and standard length of fish points out a structuring effect of size, with larger species (Gymnoscopelus spp.) occupying a higher trophic position than smaller species (Krefftichthys anderssoni and Protomyctophum spp.). Myctophids occupy an intermediate trophic position between macrozooplanktonic crustaceans and seabirds and marine mammals within the pelagic ecosystem. However, the TLs of large myctophids overlap those of crustacean-eating seabirds [e.g., Eudyptes spp. (crested penguins) and Pachyptila belcheri]. The isotopic niche of myctophids indicates that Aptenodytes patagonicus (king penguin) adults prey upon K. anderssoni when they feed for themselves, thus exemplifying the usefulness of isotopic datasets on potential prey of predators to depict trophic relationships. Myctophid (and gonostomatid) fishes are the dominant various dimensions of their ecological niche (Newsome et mesopelagic fish of the oceanic ecosystem worldwide, al. 2007). The basic isotopic concept is that an animal’s playing a key role in pelagic food webs. They mainly feed chemical composition is directly influenced by what it on planktonic crustaceans and are therefore considered to consumes. Consumers are enriched in 15N relative to their be an essential component of the tertiary level of the pelagic food, and consequently, stable-nitrogen-isotope measure- ecosystem (Pakhomov et al. 1996; Pusch et al. 2004). In ments (15N:14N, d15N) serve as indicators of a consumer turn, they are consumed by a range of predators, including trophic position. By contrast, stable-carbon signatures fish, squids, seabirds, and marine mammals (Rodhouse and (13C:12C, d13C) vary little along the food chain and, in Nigmatullin 1996; Connan et al. 2007). Myctophids thus the marine environment, d13C values are mainly used to play a pivotal role in the transfer of energy from indicate the foraging habitats of predators, including fish. zooplankton to higher trophic levels (TLs). Determining The stable isotope method is based on time-integrated their feeding habits and trophic positions are thus essential assimilated food. Hence, the isotopic signature of white for a better understanding of the functioning of the pelagic muscle was considered to be representative of the isotopic ecosystem. In the Southern Ocean (south of the Subtropical niche of fish during the months preceding sampling Front), myctophids form by far the bulk of the mesopelagic (Herzka 2005), thus contrasting with the snapshot method fish biomass; their standing stock is estimated to be 70–200 of analyzing stomach contents and avoiding biases induced 109 kg (Sabourenkov 1992; Kozlov 1995). Hulley (1990) by feeding in nets. The stable isotope method also allows lists , 43 species in the area, of which 11 are abundant in quantifying trophic levels of consumers and thus compar- the epipelagic and mesopelagic layers, where their role as ing taxonomically related and unrelated species within a prey is demonstrated by their importance in the diet of top given ecosystem (Hobson et al. 1994; Pinnegar et al. 2002; predators (Sabourenkov 1992; Kozlov 1995; Guinet et al. Cherel et al. 2008). 1996). Despite the biological significance of Southern To date, no isotopic investigations of myctophid assem- Ocean myctophids, data on their ecology is remarkably blages are available in the scientific literature, and isotopic sparse. Dietary information is restricted to the most information on myctophids from the Southern Ocean is common species and to a few assemblages with limited restricted to a few species and individuals (Rau et al. 1992; spatio-temporal scales and, in some cases, with very small Pakhomov et al. 2006). The primary objective of this study sample sizes (Pakhomov et al. 1996; Gaskett et al. 2001; was to define and compare the isotopic niches and trophic Schreeve et al. 2009). levels of coexisting myctophid fishes living in the Southern Measuring the isotopic niche of animals is a powerful Ocean. We focused on resource partitioning within the fish alternative to the conventional means of investigating assemblage occurring in the vicinity of the Polar Front, where myctophids concentrate and are targeted by top *Corresponding author: [email protected] consumers (Bost et al. 2009). Stable isotope analyses were 324 Niches of Southern Ocean myctophid fish 325 Table 1. Sizes, isotopic niches, and estimated trophic levels of myctophid fishes from Kerguelen waters. Age was estimated using standard length (Hulley 1990). Values are mean 6 SD. Species Age n SL (mm) Mass (g) d13C(%) d15N(%) C : N Trophic level Electrona antarctica Adults and 12 7168 4.661.5 221.460.5 8.960.3 3.2660.04 3.860.1 subadults E. carlsbergi Adults 12 9064 11.462.2 221.660.4 9.560.2 3.3160.03 3.960.1 E. subaspera Mostly subadults 14 99614 16.967.5 220.260.4 7.360.3 3.3160.04 3.360.1 Gymnoscopelus bolini Subadults 12 2056997614 220.560.4 9.960.5 3.2460.04 4.160.2 G. braueri Adults and 12 10767 11.562.5 222.360.7 9.860.3 3.2660.03 4.060.1 subadults G. fraseri Adults 12 8364 6.460.9 221.160.4 9.060.4 3.2660.03 3.860.1 G. nicholsi Subadults 12 12663 20.561.6 221.160.3 10.260.5 3.3360.04 4.260.2 G. piabilis Adults 12 13766 30.064.6 219.860.3 8.860.2 3.2460.02 3.760.1 Krefftichthys Adults and 12 5164 1.560.4 222.360.2 7.660.2 3.4160.05 3.360.1 anderssoni subadults Protomyctophum Mostly adults 7 5164 1.960.4 220.960.3 8.760.4 3.3060.05 3.760.1 andriashevi P. bolini Adults 12 5664 2.260.3 222.460.6 9.260.4 3.2660.03 3.960.1 P. choriodon Subadults 12 6264 3.760.7 220.060.5 7.860.3 3.4660.22 3.460.1 P. gemmatum Adults 4 7764 6.561.4 222.160.1 8.760.4 3.2060.02 3.760.1 P. tenisoni Adults 11 5261 1.760.2 222.160.3 8.160.3 3.2860.04 3.560.1 also performed on potential prey (macrozooplankton) and their vertical distribution and diet, but, instead, to collect predators (seabirds) to create a basis for the interpretation the main mesopelagic fish and macrozooplankton species of the isotopic signatures and trophic levels of myctophids occurring in Kerguelen waters in order to build biochemical within the oceanic pelagic ecosystem. reference datasets (i.e., stable isotopes and lipids) to better The present work is a part of a larger program examin- investigate trophic relationships within the pelagic ecosys- ing isotopically the trophic web at Kerguelen Islands to tem. Following Duhamel et al. (2000), the study area better interpret the isotopic niches, and thus the feeding (49u059–49u209S, 71u159–72u159E) was located in slope ecology of seabirds and marine mammals in an area where waters, where the Polar Front runs northward along and Euphausia superba (Antarctic krill) do not occur. This off the eastern shelf of the archipelago. Biological data were makes the archipelago an ideal location to investigate collected on board the 25-m-long trawler La Curieuse interspecies resource partitioning in the absence of the during two consecutive nights (from 22 to 24 January masking effect of the superabundance of a single prey on 2005). Fish and invertebrates were sampled using an segregating mechanisms. The selected sampling sites for International Young Gadoid Pelagic Trawl (IYGPT; myctophids were included within the foraging areas of opening: 12 m 3 7 m) with a 10-mm cod-end mesh Aptenodytes patagonicus (king penguins) and female (Duhamel et al. 2000). Fishing depth was estimated during Arctocephalus gazella (Antarctic fur seals) at a time of the trawling operations using relationship between meters- year (summer) when both species are known to prey on wire-out (the number of meters the wire of the collection them (Guinet et al. 2001; Bost et al. 2009). The usefulness gear was out from the collecting vessel) and depth, and it of the approach was tested first by comparing the isotopic was logged using a time-depths recorder (Mk7, Wildlife signature of myctophids with those of the food of A. Computers). Eight trawls (four per night) were performed patagonicus chicks and of the chicks themselves. Second, at different depths ranging from 50 m to 425 m. The net the comparison was extended to breeding and molting was deployed using standardized 30-min duration of adult A.
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