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Hydroacoustic surveys using hull-mounted down-looking transducers are useful for estimating pelagic fish densities; however, this method may miss shallow fish owing to the acoustic surface dead zone and vessel avoidance. Our objective was to compare pelagic fish density estimates acquired by a traditional down-looking acoustic survey to estimates obtained by a new multi-directional-towed sled capable of sampling the entire water column using upward-, sideways-, and downward-aimed transducers simultaneously. We deployed both systems concurrently in the western arm of Lake Superior during a period of stable stratification. We found the two survey approaches provided significantly different estimates of fish density in the upper water column layer (~4–9 m below the lake surface) with the sled up-looking transducer providing 56 times higher densities compared to the traditional ship down-looking method. Densities also varied significantly in the 9–14 m layer where densities were 6.2 times higher in the sled survey. Midwater trawl sampling indicated that cisco (Coregonus artedi) and rainbow smelt (Osmerus mordax) were the predominant species occupying the uppermost 14 m of the water column. The two acoustic approaches provided similar results at water column depths >14 m where rainbow smelt and kiyi (Coregonus kiyi) were predominant. Overall, the sled-based method estimates were, on average, 2.5 times higher for the whole water column. Our findings show that the new sled can reduce bias by better sampling the surface dead zone leading to more accurate estimation of pelagic fish densities for both management and research.

Groups of sympatric taxa with low inter-specific genetic differentiation, but considerable ecological differences, offer great opportunities to study the dynamics of divergence and speciation. This is the case of ciscoes (Coregonus spp.) in the Laurentian Great Lakes, which are characterized by a complex evolutionary history and are commonly described as having undergone an adaptive radiation. In this study, morphometrics, stable isotopes and transcriptome sequencing were used to study the relationships within the Coregonus artedi complex in western Lake Superior. We observed general concordance for morphological, ecological and genomic variation, but the latter was more taxonomically informative as it showed less overlap among species in multivariate space. Low levels of genetic differentiation were observed between individuals morphologically identified as C. hoyi and C. zenithicus, and we hypothesize this could be associated with recent hybridization between the two species. Transcriptome-based single nucleotide polymorphisms exhibited significant divergence for genes associated with vision, development, metabolism and immunity, among species that occupy different habitats. This study highlights the importance of using an integrative approach when studying groups of taxa with a complex evolutionary history, as individual-level analyses of multiple independent datasets can provide a clearer picture of the patterns and processes associated with the origins of biodiversity.

Local adaptation can drive diversification of closely related species across environmental gradients and promote convergence of distantly related taxa that experience similar conditions. We examined a potential case of adaptation to novel visual environments in a species flock (Great Lakes salmonids, genus Coregonus) using a new amplicon genotyping protocol on the Oxford Nanopore Flongle and MinION. We sequenced five visual opsin genes for individuals of Coregonus artedi, Coregonus hoyi, Coregonus kiyi, and Coregonus zenithicus. Comparisons revealed species-specific differences in a key spectral tuning amino acid in rhodopsin (Tyr261Phe substitution), suggesting local adaptation of C. kiyi to the blue-shifted depths of Lake Superior. Ancestral state reconstruction demonstrates that parallel evolution and “toggling” at this amino acid residue has occurred several times across the fish tree of life, resulting in identical changes to the visual systems of distantly related taxa across replicated environmental gradients. Our results suggest that ecological differences and local adaptation to distinct visual environments are strong drivers of both evolutionary parallelism and diversification.

Stock assessments are critical to modern fisheries management, supporting the calculation of key reference variables used to make informed management decisions. However, there is still considerable uncertainty as to which class of assessment models is appropriate to use under different circumstances. A common class of models used when age data are available are statistical catch-at-age assessment (SCAA) models, which track annual cohorts through time. When age data are unavailable, as is often the case in invertebrate fisheries where the lack of a bony structure such as otoliths makes aging difficult, statistical catch-at-size assessment (SCSA) models are more often employed, tracking fish or invertebrates through time by size-classes rather than ages. Do SCAA models actually perform better than SCSA models when age data are available, or is this just an assumption we make in fisheries research and management? We examined this question by evaluating the effectiveness of both SCAA and SCSA models in characterizing cisco, Coregonus artedi, population dynamics in Thunder Bay, Ontario. Both models were fit using an integrated framework with multiple sources of data including hydroacoustic estimates of spawning stock, fishery-dependent and -independent age/length compositions, and harvest data. Our results suggest that for cisco in Thunder Bay, data-limitations related to lack of size-composition data over the size range for which cisco growth is rapid resulted in difficulty estimating relative year-class strength within a SCSA. This led to parameter confounding and ultimately the inability to estimate natural mortality within a SCSA. This hampered the utility of a SCSA model in comparison with a SCAA model when age-composition data were available.

We examined the spatial scale of recruitment variability for disparate cisco (Coregonus artedi) populations in the Great Lakes (n = 8) and Minnesota inland lakes (n = 4). We found that the scale of synchrony was approximately 400 km when all available data were utilized; much greater than the 50-km scale suggested for freshwater fish populations in an earlier global analysis. The presence of recruitment synchrony between Great Lakes and inland lake cisco populations supports the hypothesis that synchronicity is driven by climate and not dispersal. We also found synchrony in larval densities among three Lake Superior populations separated by 25–275 km, which further supports the hypothesis that broad-scale climatic factors are the cause of spatial synchrony. Among several candidate climate variables measured during the period of larval cisco emergence, maximum wind speeds exhibited the most similar spatial scale of synchrony to that observed for cisco. Other factors, such as average water temperatures, exhibited synchrony on broader spatial scales, which suggests they could also be contributing to recruitment synchrony. Our results provide evidence that abiotic factors can induce synchronous patterns of recruitment for populations of cisco inhabiting waters across a broad geographic range, and show that broad-scale synchrony of recruitment can occur in freshwater fish populations as well as those from marine systems.

International audience; Fish species diversity can be lost through interacting stressors including habitat loss, stocking and overfishing. Although a multitude of stressors have played a role in the global decline of coregonid (Coregonus spp.) diversity, a number of contemporary studies have identified habitat loss stem-ming from eutrophication as the primary cause. Unfortunately, reconstructing the role of fishing and stocking practices can be difficult, because these records are incomplete or appear only in hard-to-access historic grey literature. Based on an illustrative set of historic and contemporary studies, we describe how fisheries management practices may have contributed to coregonid di-versity loss in European and North American lakes. We provide case studies examining how fish-ing and stocking may reduce coregonid diversity through demographic decline and introgressive hybridization. In some lakes, fisheries management practices may have led to a loss of coregonid diversity well before issues with habitat degradation manifested. Our review suggests that fish conservation policies could beneficially consider the relative importance of all stressors, including management practices, as potential drivers of diversity loss.

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September–November) and two northern Lake Michigan transects (April, July, and September). In Lake Superior, consumption (primarily by cisco Coregonus artedi ) exceeded Bythotrephes production at all offshore sites in September–November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site during September–October. In Lake Michigan, consumption by fishes (primarily alewife Alosa pseudoharengus ) exceeded production at nearshore sites (up to 178%), but not in deeper sites ( Bythotrephes never subsequently declined. Using generalized additive models, temperature, and not fish consumption nor zooplankton prey density, best explained variability in Bythotrephes biomass. The non-linear pattern revealed Bythotrephes to increase with temperature up to 16 °C, and then decline between 16 and 23 °C. We discuss how temperature likely has direct negative impacts on Bythotrephes when temperatures near 23 °C, but speculate that predation also contributes to declining biomass when temperatures exceed 16 °C.

The relative importance of predation and food availability as contributors to larval cisco ( Coregonus artedi ) mortality in Lake Superior were investigated using a visual foraging model to evaluate potential predation pressure by rainbow smelt ( Osmerus mordax ) and a bioenergetic model to evaluate potential starvation risk. The models were informed by observations of rainbow smelt, larval cisco, and zooplankton abundance at three Lake Superior locations during the period of spring larval cisco emergence and surface-oriented foraging. Predation risk was highest at Black Bay, ON, where average rainbow smelt densities in the uppermost 10 m of the water column were >1000 ha −1 . Turbid conditions at the Twin Ports, WI-MN, affected larval cisco predation risk because rainbow smelt remained suspended in the upper water column during daylight, placing them alongside larval cisco during both day and night hours. Predation risk was low at Cornucopia, WI, owing to low smelt densities ( −1 ) and deep light penetration, which kept rainbow smelt near the lakebed and far from larvae during daylight. In situ zooplankton density estimates were low compared to the values used to develop the larval coregonid bioenergetics model, leading to predictions of negative growth rates for 10 mm larvae at all three locations. The model predicted that 15 mm larvae were capable of attaining positive growth at Cornucopia and the Twin Ports where low water temperatures (2–6 °C) decreased their metabolic costs. Larval prey resources were highest at Black Bay but warmer water temperatures there offset the benefit of increased prey availability. A sensitivity analysis performed on the rainbow smelt visual foraging model showed that it was relatively insensitive, while the coregonid bioenergetics model showed that the absolute growth rate predictions were highly sensitive to input parameters (i.e., 20% parameter perturbation led to order of magnitude differences in model estimates). Our modelling indicated that rainbow smelt predation may limit larval cisco survival at Black Bay and to a lesser extent at Twin Ports, and that starvation may be a major source of mortality at all three locations. The framework we describe has the potential to further our understanding of the relative importance of starvation and predation on larval fish survivorship, provided information on prey resources available to larvae are measured at sufficiently fine spatial scales and the models provide a realistic depiction of the dynamic processes that the larvae experience.

SUMMARY 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus artedi) provides a resource subsidy, in the form of energy-rich cisco eggs, from the offshore pelagic to the nearshore benthic community over winter, when alternate prey production is likely to be low. 2. We tested this hypothesis using fish and macroinvertebrate surveys, fish population demographics, diet and stable isotope analyses, and bioenergetics modelling. 3. The benthic, congeneric lake whitefish (C. clupeaformis) was a clear beneficiary of cisco spawning. Cisco eggs represented 16% of lake whitefish annual consumption in terms of biomass, but 34% of energy (because of their high energy density: >10 kJ g wet mass 1 ). Stable isotope analyses were consistent with these results and suggest that other nearshore fish species may also rely on cisco eggs. 4. The lipid content of lake whitefish liver almost doubled from 26 to 49% between November and March, while that of muscle increased from 14 to 26% over the same period, suggesting lake whitefish were building, rather than depleting, lipid reserves during winter. 5. In the other Laurentian Great Lakes, where cisco populations remain very low and rehabilitation efforts are underway, the offshore-to-nearshore ecological link apparent in Lake Superior has been replaced by non-native planktivorous species. These non-native species spawn in spring have smaller eggs and shorter incubation periods. The rehabilitation of cisco in these systems should reinstate the onshore subsidy as it has in Lake Superior.

Acoustic methods are used to estimate the density of pelagic fish in large lakes with results of midwater trawling used to assign species composition. Apportionment in lakes having mixed species can be challenging because only a small fraction of the water sampled acoustically is sampled with trawl gear. Here we describe a new method where single echo detections (SEDs) are assigned to species based on classification tree models developed from catch data that separate species based on fish size and the spatial habitats they occupy. During the summer of 2011, we conducted a spatially-balanced lake-wide acoustic and midwater trawl survey of Lake Superior. A total of 51 sites in four bathymetric depth strata (0–30 m, 30–100 m, 100–200 m, and >200 m) were sampled. We developed classification tree models for each stratum and found fish length was the most important variable for separating species. To apply these trees to the acoustic data, we needed to identify a target strength to length (TS-to-L) relationship appropriate for all abundant Lake Superior pelagic species. We tested performance of 7 general (i.e., multi-species) relationships derived from three published studies. The best-performing relationship was identified by comparing predicted and observed catch compositions using a second independent Lake Superior data set. Once identified, the relationship was used to predict lengths of SEDs from the lake-wide survey, and the classification tree models were used to assign each SED to a species. Exotic rainbow smelt (Osmerus mordax) were the most common species at bathymetric depths 100 m (384 million; 6.0 kt). Cisco (Coregonus artedi) were widely distributed over all strata with their population estimated at 182 million (44 kt). The apportionment method we describe should be transferable to other large lakes provided fish are not tightly aggregated, and an appropriate TS-to-L relationship for abundant pelagic fish species can be determined.

Diel patterns of distribution of fishes in nearshore (15–80 m depth) and offshore (>80 m) waters of the Apostle Islands region of Lake Superior were described using bottom trawls, mid-water trawls, and acoustic gear during day and night sampling. These data revealed three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no migration. DVM was expressed by fishes migrating from benthopelagic to pelagic strata and DBM was expressed by fishes migrating horizontally from deeper waters in the day to shallower waters at night while remaining within the benthopelagic stratum. Most fishes that did not exhibit diel migration showed increased nighttime densities as a result of increased activity and movement from benthic to benthopelagic strata. Rainbow Smelt (Osmerus mordax), Cisco (Coregonus artedi), Bloater (C. hoyi), Kiyi (C. kiyi), juvenile Trout-Perch (Percopsis omiscomaycus), and adult siscowet (Salvelinus namaycush siscowet) exhibited DVM. Lake Whitefish (C. clupeaformis), lean Lake Trout (Salvelinus namaycush namaycush), and juvenile siscowet exhibited DBM. Adult Trout-Perch and adult Pygmy Whitefish (Prosopium coulteri) exhibited a mixture of DBM and DVM. Burbot (Lota lota), Slimy Sculpin (Cottus cognatus), Spoonhead Sculpin (C. ricei), and Deepwater Sculpin (Myoxocephalus thompsonii) did not exhibit diel migration, but showed evidence of increased nocturnal activity. Ninespine Stickleback (Pungitius pungitius) exhibited a mixture of DVM and non-migration. Juvenile Pygmy Whitefish did not show a diel change in density or depth distribution. Species showing ontogenetic shifts in depth distribution with larger, adult life stages occupying deeper waters included, Rainbow Smelt, lean and siscowet Lake Trout, Lake Whitefish, Pygmy Whitefish, Ninespine Stickleback and Trout-Perch. Of these species, siscowet also showed an ontogenetic shift from primarily DBM as juveniles to primarily DVM as adults. Across all depths, fishes expressing DVM accounted for 73% of the total estimated community areal biomass (kg ha−1) while those expressing DBM accounted for 25% and non-migratory species represented 2% of the biomass. The proportion of total community biomass exhibiting DVM increased with depth, from 59% to 95% across ≤30 m to >90 m depth zones. Along the same depth gradient, the proportion of total community biomass exhibiting DBM declined from 40% to 1%, while non-migrators increased from 1% to 4%. These results indicate that DVM and DBM behaviors are pervasive in the Lake Superior fish community and potentially provide strong linkages that effect coupling of benthic and pelagic and nearshore and offshore habitats.

Detailed diet linkages within the offshore (> 80 m bathymetric depth) food web of Lake Superior are currently not well identified. We used analyses of fish stomach contents to create an empirically based food web model of the Lake Superior offshore fish community. Stomachs were collected seasonally (spring, summer, and fall) from nine offshore locations in 2005, using bottom and midwater trawls. In total, 2643 stomachs representing 12 fish species were examined. The predominant fish species collected were deepwater sculpin ( Myoxocephalus thompsonii ), siscowet ( Salvelinus namaycush siscowet ), kiyi ( Coregonus kiyi ), and cisco ( Coregonus artedi ). Mysis diluviana was the most common prey item, indicating that changes in Mysis abundance could have a profound impact on the entire offshore food web. Mysis was the primary diet item of deepwater sculpin (≥ 53% by mass) and kiyi (≥ 96% by mass) regardless of depth or season. The invasive Bythotrephes was an important diet component of the pelagic cisco in summer and fall. Deepwater sculpin were the primary diet item of siscowet (≥ 52% by mass), with coregonines appearing in the diet of larger (> 400 mm) siscowet. Non-metric multidimensional scaling analysis indicated that there were no statistically significant seasonal or site-specific differences in diets of deepwater sculpin, cisco, or kiyi. Site was the primary structuring factor in siscowet diets. Generally, in Lake Superior, the diet items of the dominant offshore species did not appear to be in danger from those types of major ecological shifts occurring in the lower Laurentian Great Lakes.

Recent studies on Lake Superior suggest that diel vertical migration (DVM) of prey (generalized Coregonus spp.) may be influenced by the density of predatory siscowet (Salvelinus namaycush). We empirically evaluated this hy- pothesis using data from acoustic, midwater trawl, and bottom trawl sampling at eight Lake Superior sites during three seasons in 2005 and a subset of sites in 2006. We expected the larger-bodied cisco (Coregonus artedi) to exhibit a shal- lower DVM compared with the smaller-bodied kiyi (Coregonus kiyi). Although DVM of kiyi and cisco were consistent with expectations of DVM as a size-dependent, predator-mediated process, we found no relationship between siscowet density and the magnitude of DVM of either coregonid. Cisco appear to have a size refuge from siscowet predation. Kiyi and siscowet co-occur in demersal habitat > 150 m during the day, where visual predation is unlikely, suggesting predator avoidance is not a factor in the daytime distribution of kiyi. Seasonal patterns of kiyi DVM were consistent with reported DVM of their primary prey Mysis relicta. Our results suggest that consideration of nonvisual foraging, rather than light- based foraging theory (i.e., the antipredation window), is necessary to understand the processes driving DVM in deepwater systems. Resume´ : Des etudes recentes sur le lac Superieur laissent croire que la migration verticale journaliere (DVM) des proies (les Coregonus spp. en general) est influencee par la densitedu predateur, le touladi siscowet (Salvelinus namaycush). Nous evaluons cette hypothese de maniere empirique a l'aide de donnees provenant d'echantillonnages acoustiques et de prelevements aux chaluts pelagique et benthique dans huit sites au lac Superieur durant trois saisons en 2005 et dans un sous-ensemble de sites en 2006. Nous nous attendions ace que le cisco de lac (Coregonus artedi) de plus grande taille ait

Evidence from small lakes suggests that predation on larval ciscoes Coregonus artedi by nonnative rainbow smelt Osmerus mordax can lead to cisco suppression or extirpation. However, evidence from larger lakes has led to equivocal conclusions. In this study, we examine the potential predation effects of rainbow smelt in two adjacent but contrasting embayments in Lake Superior (Thunder and Black bays, Ontario). During May 2006, we sampled the ichthyoplankton, pelagic fish communities, and diet composition of rainbow smelt in both bays. Using acoustics and midwater trawling, we estimated rainbow smelt densities to be 476 ± 34/ha (mean ± SE) in Thunder Bay and 3,435 ± 460/ha in Black Bay. We used a bioenergetics model to estimate the proportion of cisco larvae consumed by rainbow smelt. Our results suggest that predation by rainbow smelt accounts for 15–52% and 37–100% of the mortality of larval ciscoes in Thunder and Black bays, respectively, depending on the predator feeding rate and the scale of p...

Populations of cisco Coregonus artedi in the Laurentian Great Lakes supported large-scale commercial fisheries and were the primary forage of piscivores during the first half of the 20th century. However, by 1970 populations had collapsed in all of the lakes. Since then, ciscoes have staged a recovery in Lake Superior. In this synthesis, we describe the status of ciscoes in Lake Superior during 1970–2006 and provide a comprehensive review of their ecology. Better understanding of age estimation techniques, application of hydroacoustic and midwater trawl sampling, and compilation of long-term data sets have advanced our understanding of the species. Management agencies contemplating rehabilitation of cisco populations should recognize that (1) knowledge of cisco ecology and population dynamics is increasing; (2) ciscoes are long-lived; (3) Great Lakes populations are probably composed of both shallow-water and deepwater spawning forms; (4) large year-classes can be produced from small adult stocks...

Management efforts to rehabilitate lake trout Salvelinus namaycush in Lake Superior have been successful and the recent increase in their numbers has led to interest in measuring biomass of pelagic prey fish species important to these predators. Lake Superior cisco Coregonus artedi currently support roe fisheries and determining the sustainability of these fisheries is an important management issue. We conducted acoustic and midwater trawl surveys of the western arm of Lake Superior during three periods: summer (July–August), October, and November 2006 to determine if a single survey can be timed to estimate biomass of both prey fish and spawning cisco. We evaluated our methods by comparing observed trawl catches of small (

To improve our ability to assess larval cisco (Coregonus artedi) populations in Lake Superior, we conducted a study to compare several sampling strategies. First, we compared density estimates of larval cisco concurrently captured in surface waters with a 2 × 1-m paired neuston net and a 0.5-m (diameter) conical net. Density estimates obtained from the two gear types were not significantly different, suggesting that the conical net is a reasonable alternative to the more cumbersome and costly neuston net. Next, we assessed the effect of tow pattern (sinusoidal versus straight tows) to examine if propeller wash affected larval density. We found no effect of propeller wash on the catchability of larval cisco. Given the availability of global positioning systems, we recommend sampling larval cisco using straight tows to simplify protocols and facilitate straightforward measurements of volume filtered. Finally, we investigated potential trends in larval cisco density estimates by sampling four time periods during the light period of a day at individual sites. Our results indicate no significant trends in larval density estimates during the day. We conclude estimates of larval cisco density across space are not confounded by time at a daily timescale. Well-designed, cost effective surveys of larval cisco abundance will help to further our understanding of this important Great Lakes forage species.

Systematic underestimation of fish age can impede understanding of recruitment variability and adaptive strategies (like longevity) and can bias estimates of survivorship. We suspected that previous estimates of annual survival (S; range = 0.20-0.44) for Lake Superior ciscoes Coregonus artedi developed from scale ages were biased low. To test this hypothesis, we estimated the total instantaneous mortality rate of adult ciscoes from the Thunder Bay, Ontario, stock by use of cohort-based catch curves developed from commercial gill-net catches and otolith-aged fish. Mean S based on otolith ages was greater for adult females (0.80) than for adult males (0.75), but these differences were not significant. Applying the results of a study of agreement between scale and otolith ages, we modeled a scale age for each otolith-aged fish to reconstruct catch curves. Using modeled scale ages, estimates of S (0.42 for females, 0.36 for males) were comparable with those reported in past studies. We conducted a No...

We compared density estimates from day bottom trawl tows against night midwater trawl tows and acoustic gear to test the hypothesis that adult lake herring (≥250 mm) are underestimated by day bottom trawl tows during the annual USGS spring fish community survey in Lake Superior. We found average density at nine nearshore stations was significantly higher at night (21.3 adult fish/ha) compared to day (1.0 adult fish/ha; p = 0.0119). At nine offshore stations, no lake herring were captured during the day but density averaged 39.6 adult fish/ha at night. At a lakewide scale (n = 18 stations), precision (relative standard error) was much better using night midwater trawls and acoustic gear (37%) compared to day bottom trawls (100%). Moderate sample size increases using the former methodology would likely bring precision within recommended levels (≤30%) for stock-recruit data sets. Our results suggest that 1) population abundances of adult lake herring in Lake Superior are much higher than previously ...

Historically, commercial fishers harvested Lake Superior lake herring Coregonus artedi for their flesh, but recently operators have targeted lake herring for roe. Because no surveys have estimated spawning female abundance, direct estimates of fishing mortality are lacking. The primary objective of this study was to determine the feasibility of using acoustic techniques in combination with midwater trawling to estimate spawning female lake herring densities in a Lake Superior statistical grid (i.e., a 10′ latitude × 10′ longitude area over which annual commercial harvest statistics are compiled). Midwater trawling showed that mature female lake herring were largely pelagic during the night in late November, accounting for 94.5% of all fish caught exceeding 250 mm total length. When calculating acoustic estimates of mature female lake herring, we excluded backscattering from smaller pelagic fishes like immature lake herring and rainbow smelt Osmerus mordax by applying an empirically derived thresh...

We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female age-7 ciscoes (Coregonus artedi) captured from a spawning aggregation in Thunder Bay, Lake Superior, during November 2010. We also determined PCB concentrations in the ovaries and somatic tissue of five additional female ciscoes (ages 5-22). All 55 of these ciscoes were in ripe or nearly ripe condition. Bioenergetics modeling was used to determine the contribution of the growth dilution effect toward a difference in PCB concentrations between the sexes, as females grew substantially faster than males. Results showed that the PCB concentration of males (mean = 141 ng/g) was 43% greater than that of females (mean = 98 ng/g), and this difference was highly significant (P