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Spatial Structure and Movement of Blue Cod Parapercis Colias in Doubtful Sound, New Zealand, Inferred from Δ13c and Δ15n

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Spatial Structure and Movement of Blue Cod Parapercis Colias in Doubtful Sound, New Zealand, Inferred from Δ13c and Δ15n Vol. 359: 239–248, 2008 MARINE ECOLOGY PROGRESS SERIES Published May 5 doi: 10.3354/meps07349 Mar Ecol Prog Ser Spatial structure and movement of blue cod Parapercis colias in Doubtful Sound, New Zealand, inferred from δ13C and δ15N Kirsten L. Rodgers, Stephen R. Wing* Department of Marine Science, University of Otago, 310 Castle Street, Dunedin 9054, New Zealand ABSTRACT: Migrations and home range shifts by adult blue cod Parapercis colias provide an impor- tant means of dispersal among resident subpopulations within the Doubtful–Thompson Sound com- plex, a fjord system in southwestern New Zealand. In the present study we used stable isotope analy- sis (δ13C and δ15N) to investigate regional differences in the carbon sources for, and likely migration patterns among, subpopulations of P. colias. We found consistent differences in δ13C and δ15N of blood and muscle collected from outer coast (mean ± SE: blood, δ13C = –18.4 ± 0.2, δ15N = 11.5 ± 0.2; muscle, δ13C = –17.9 ± 0.1, δ15N = 13.0 ± 0.1) and inner fjord (mean ± SE: blood, δ13C = –19.9 ± 0.5, δ15N = 9.5 ± 0.3; muscle, δ13C = –19.3 ± 0.4, δ15N = 11.6 ± 0.2) locations, likely reflecting long-term (>1 yr) residency. Inner fjord fish had evidence of significant input of recycled carbon from chemoau- totrophic bacteria. On the outer coast, δ15N and δ13C reflected a diet likely supported primarily by new production from macroalgae and phytoplankton. To provide information on isotopic turnover rate of blood and muscle we translocated fish from the outer coast to an inner fjord site and measured change in δ15N and δ13C after 160 d. These data showed a depletion of 13C and 15N with shift in habi- tat and provided a basis for classifying likely migrants (<160 d) to each study site. The conclusion that populations of blue cod within Doubtful Sound are made up of long-term residents with some subsidy from the outer coast has implications for the efficacy of 2 newly established marine reserves in Doubt- ful Sound and others in Fiordland. KEY WORDS: Stable isotopes · Metapopulation · Connectivity · Diet · Isotopic turnover · Temperate reef fish Resale or republication not permitted without written consent of the publisher INTRODUCTION ing, home range shifts or migrations may also substan- tially contribute to population connectivity (Thorrold et Information on metapopulation structure, in particu- al. 2001). The relative rates and spatial scale of these lar connectivity among local populations, remains a exchanges among local subpopulations significantly fundamental theme in marine population ecology influence rapid population replenishment and local (Kritzer & Sale 2006) essential for understanding buildup of reproductive potential and therefore have spatial population dynamics (Pulliam 1988, Hixon et large implications for population dynamics and the al. 2002) and therefore the effectiveness of marine efficacy of spatial management of marine species. protected areas and other spatial management In some situations information on these types of approaches (Quinn et al. 1993). Connectivity by ex- migrations can be obtained from stable isotope signa- change of larvae, recruits, or juveniles is an important tures (Hobson 1999). The stable isotopic composition of mechanism for determining the spatial and demo- a consumer provides an integrated reflection of the iso- graphic structure of coastal marine species (Rough- topic composition of its diet (DeNiro & Epstein 1978, garden et al. 1985, Palumbi 2003). However, for some 1981, Tieszen et al. 1983) with a predictable amount species movement of adults in the form of natal hom- of enrichment at each trophic level (Rau et al. 1983, *Corresponding author. Email: [email protected] © Inter-Research 2008 · www.int-res.com 240 Mar Ecol Prog Ser 359: 239–248, 2008 McCutchan et al. 2003). If animals move between lation networks within fjords, with strong reproductive habitats with isotopically distinct prey communities, source–sink structures (S. R. Wing et al. 2003). The iso- these signatures can be used to infer geographic lated basins of the inner fjords and predominant estu- origins, differentiate populations utilizing different arine circulation play an important role in limiting lar- habitats (Dufour et al. 1998, Melville & Connolly 2003) val dispersal and gene flow among fjords for marine and trace movement patterns (Fry 1981, 1983, 2006, organisms with a larval dispersal phase. Evidence for Kurle & Worthy 2002, Fry et al. 2003). The isotopic this comes from several sources including genetic composition of a consumer will gradually change to studies of the eleven-armed sea star Coscinasterias reflect the new diet, thus serving as a natural marker muricata (Sköld et al. 2003, Perrin et al. 2004), the sea for distinguishing recent immigrants from those that urchin Evechinus chloroticus (Perrin et al. 2003), and are equilibrated to the local isotopic signature (Fry brachiopods Liothyrella neozelanica and Terebratella 1983, Herzka 2005, Guelinckx et al. 2007). Accord- sanguinea (Ostrow et al. 2001). ingly, an understanding of the isotopic patterns in the We characterized spatial variability in the composi- environment is required to distinguish population tion of diet and resulting δ13C and δ15N of blood and structure and trace animal movement (Rubenstein & muscle in blue cod Parapercis colias collected from dif- Hobson 2004). ferent regions of the Doubtful–Thompson Sound com- The present study used δ13C and δ15N of blood and plex. We then compared site-specific δ13C and δ15N of muscle tissue to investigate spatial structure of blue P. colias with the primary carbon source pools (macro- cod Parapercis colias populations in the Doubtful– algae, phytoplankton and chemoautotrophic endosym- Thompson Sound complex, a fjord system on the bionts). To simulate migration we investigated isotopic southwest coast of New Zealand’s South Island. P. col- turnover in blood and muscle tissue in situ by relocat- ias are a conspicuous benthic predator in the rocky ing individuals from the outer to inner region of Doubt- reef fish assemblage of southern New Zealand and ful Sound and quantifying turnover in δ13C and δ15N support significant commercial and recreational fish- between time of relocation and recapture. Stomach eries around the South Island. Little is known about contents (immediate past) as well as δ13C and δ15N of spawning behavior or early life history, including the blood (recent past) and muscle (long-term past) pro- distribution of juveniles. Growth rates can vary within vided an indication of individual dietary history inte- and among areas, with a maximum age of 23 yr (S. R. grated over a range of timescales (Tieszen et al. 1983). Wing unpubl. data). Adults are typically relatively The measure of turnover in these tissues was then used sedentary with restricted home ranges (Mace & John- as a basis for identifying migrants between the outer son 1983, Cole et al. 2000). As a consequence P. colias coast and inner fjord habitats. These data provided are considered to be vulnerable to localized depletion valuable insight into the pattern of movement and because of a highly divided stock structure and moder- home range shifts for P. colias within the fjord, which ate rates of movement among habitats (Carbines & are discussed in the context of their implications for McKenzie 2004). source–sink structure and population replenishment These concerns are particularly important in the 14 within 2 recently established marine reserves in the fjords that indent the southwest coast of the South region. Island, New Zealand, which present a highly frag- mented and diverse array of habitats. The marine habi- tats in Fiordland also present a range of carbon sources MATERIALS AND METHODS with chemoautotrophic, terrestrial and freshwater pri- mary producers particularly important in the inner Sample collection. Adult blue cod were sampled fjord habitats (Lusseau & Wing 2006, McLeod & Wing from 2 inner fjord (n = 35) and 2 outer coast (n = 34) 2007). In contrast, the outer regions of the fjords are locations in the Doubtful–Thompson Sound complex exposed to the open ocean and have productive and (45° 30’ S, 167° 0’ E) between May 2004 and January diverse phytoplankton and macroalgal-based commu- 2005 (Fig. 1), chosen to represent distinct regions in the nities (Wing et al. 2007). The result is that food web isotopic signatures of primary producers present in the structure and δ15N and δ13C of primary producers in fjord (Cornelisen et al. 2007). Fish were immediately the inner fjords are distinct from that on the outer frozen and later thawed at Portobello Marine Labora- coast, indicating that migrants among habitats can be tory (PML), where samples of blood and muscle tissue distinguished isotopically (Cornelisen et al. 2007, Wing were collected. Stomachs were removed and pre- et al. 2007). served in 70% isopropyl alcohol (IPA). One consequence of the fragmented habitat struc- Sample preparation for stable isotope analysis ture within Fiordland is that marine populations may (SIA). Blood and muscle tissue samples were dried at be less well mixed and more likely to persist as popu- 60°C for 2 d and ground to a fine powder by mortar and Rodgers & Wing: Spatial structure and movement of Parapercis colias 241 muscle tissue and among regions for each tissue. One- way ANOVA was used to compare δ13C and δ15N among sites, with a Tukey honestly significant differ- ence (HSD) post-hoc test used to resolve pairwise dif- ferences. For analysis at the regional level, in absence of significant differences at the site level, samples from the 2 outer fjord sites and those from the 2 inner fjord sites were pooled. Stomach content analysis. Stomach contents were sorted and classified into 8 broad taxonomic cate- gories: algae, thaliacea, crustacea, mollusca, poly- chaeta, echinodermata, fish, Solemya parkinsoni and other unidentified prey items. S. parkinsoni is a chemoautotrophic clam that has a distinctively de- pleted isotopic signature (mean ± SE: δ15N = –5.5 ± 0.5; δ13C = –31.3 ± 0.1) that may have a significant effect on the δ13C signature of animals that feed on it; therefore, it was shown as a separate category.
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